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REV. E. WASN ANN “ee 


SYCHOLOG GYorANTS 
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THE LIBRARY 
OF 
THE UNIVERSITY 
OF CALIFORNIA 
LOS ANGELES 


John M. Mehutens 





COMPARATIVE STUDIES 


IN THE 


Psychology of Ants and of 
Higher Animals. 


BY 


ERIC WASMANN, S. J. 


Plus enim formicularum et apicularum opera 
stupemus quam immensa corpora balaenarum, 
(S. Augustine, De Civit, Dei, |. 22, c. 24, n. 5.) 


Authorized English Version of the second German Edition. 
Enlarged and revised by the Author. 


St. Louis, Mo., AND FREIBURG, (BADEN), 
Published by B. HERDER. 
1905. 
LONDON AND EDINBURGH: SANDS & CO. 


COPYRIGHT 1905 
BY 
JoSEPH GUMMERSBACH. 


— BECKTOLD— 
PRINTING AND BOOK MFG. CO. 
ST. LOUIS, MO. 


PREFACE TO THE FIRST EDITION. 


OME time ago we published an essay entitled 

“Instinct and Intelligence in the Animal King- 
dom,” examining in detail the concepts of instinct 
and intelligence, with their application to animals. 
The discussion showed that intelligence is the spiritual 
power of abstraction, and not the mere faculty of 
forming complex sense-representations; for the laws 
of association in sense-perceptions belong to the sphere 
of instinctive sensitive life and not to spiritual intelli- 
gence. Now, what modern animal psychology terms 
“intelligence of animals,” is nothing but inborn 
instinct, raised to a higher level of perfection by the 
individual’s sensuous experience. This, in its turn, 
is based on the very same laws of association of sense- 
representations. Hence, there is no reason for ascrib- 
ing to animals intelligence in the strict sense. Indeed, 
our reasoning led us to take a further step, and we 
proved that animals have no intelligence at all. If 
they were gifted with a spiritual power of abstraction, 
it would necessarily be manifested in their outward 
actions, especially by the formation of an arbitrary 
phonetic or graphic language. Animals, however, 
have no language; hence, they have no intelligence. 

Besides, we have shown in the same essay that 
the manifestations of the psychic life, both of higher 
and of lower animals, are to be judged according to 
one and the same critical standard. The anatomical 

iii 


Preface to the First Edition. 


difference, that exists between the sense organs and 
the nervous system of Arthropods on the one hand and 
of Vertebrates on the other, is not a sufficient a priori 
reason for ascribing intelligence to the latter only and 
denying it to the former. The proof of these asser- 
tions forms the groundwork of the present essay. We 
shall compare more at length the psychic life of the 
most “intelligent” Arthropods, namely the ants, with 
that of the higher Vertebrates and of man. From this 
discussion we shall learn, whether the “missing link,”’ 
with which modern evolutionists hope to bridge over 
the chasm between the instinct of animals and the 
spiritual soul of man, is to be looked for in ants or 
in the higher Vertebrates, or whether, in fact, it exists 
at all. Biologists will be pleased to find that the 
present essay contains many new observations on the 
habits of ants and their guests. 


PREFACE TO THE SECOND EDITION. 


HE numerous observations of modern scientists, 

illustrating the relations between the psychic life 
of ants and of higher animals, have been extensively 
utilized in preparing this second edition. We have 
paid due regard to the observations and experiments, 
published since 1897, on the differentiation of castes 
in bee-hives. Besides, we have turned to account the 
results of a statistical chart now completed, extending 
over five years and comprising all the colonies of 
Formica sanguinea in the neighborhood of Exaten, 
Holland. This ant is the most interesting of all 
European species. Thus we are able to publish many 
new facts of interest in scientific biology regarding 
the slave-making habits of this ant, its methods of 
nest construction, its relationship to its guest 
Lomechusa, and the influence of the latter in the 
differentiation of castes in ant communities. Finally, 
two additional illustrations of Lomechusa strumosa 
and of its larva have been added in the text. 


TRANSLATOR’S PREFACE. 


TASMANN’S “Instinct and Intelligence in the 
Animal Kingdom,” which appeared recently in 
an English dress (Herder, St. Louis, Mo.), was so 
favorably received, that it has been thought advisable 
to follow it up with this translation of another essay 
by the same author. These two books supplement 
each other, as may be gathered from the frequent 
cross references they contain, and, more especially, 
from the close relationship of the subjects of which 
they treat. 

The best recommendation of Wasmann’s biological 
and psychological essays is given in the following 
lines of W. M. Wheeler, Prof. of the University of 
Texas:1 “Wasmann in his numerous writings has 
undoubtedly done much, at least in Germafiy, towards 
the exposure of this pseudo-psychology (of Brehm, 
Buechner and others) and a more rational conception 
of ant behavior. His long familiarity with these 
animals and their guests has given him a singularly 
lucid insight into their activities. My own more 
limited observations on our North American species 
lead me to agree with him so far as the facts are con- 
cerned, and many of the inferences which he has 
drawn from them.” As to his additional remark: 
“T am constrained to say, however, that I cannot adopt 


1) “The Compound and Mixed Nests of American Ants,” in 
“American Naturalist,” Vol. XXV, 1901, p. 808. 


vi 


Translator’s Preface. 


either his psychological definitions or his psychogenetic 
reservations,”—we call the critic’s attention to the end 
of the fourth chapter of “Instinct and Intelligence in 
the Animal Kingdom,” where Prof. Wheeler’s objec- 
tions have been answered. 

In order to make the English translation more 
valuable for North America, the author has kindly 
added a series of notes and observations on the ant 
fauna of the United States. He has added, moreover, 
the figures representing the North American form of 
Formica sanguinea and that of its guest Xenodusa 
cava. The present work, therefore, is more than a 
translation; it may be called a new edition, revised 
and enlarged by Father Wasmann. 


Canisius College, Buffalo, N. Y. 


vii 


CONTENTS. 





PAGE. 


Pretace to the first: edition... 22032) See er 
Preface to the second edition .........cccceccccccces 
WERORERIOES “BERTR CE eS eg se al Sans a eee tas 


INTRODUCTION. 


Some ancient and modern views of the psychic life of 
ants and of higher animals......3...:.-.ccceccses 


CHAPTER I. 
COMMUNITY LIFE IN THE ANIMAL KINGDOM. 


1. A general view of the forms of animal communities. 
Different degrees of community life in the animal kingdom. 
Communities of bees and ants compared. Ant states, the 
most perfect animal societies, both simple and complex. 
2. The social foundations of ant states............. eiese' 
Polymorphism, the organic foundation of ant societies. 
Psychic ties in ant colonies. It is untenable to identify the 
human “state”? with that of ants. ‘‘Automatic” instincts 
cannot sufficiently account for the latter. 
3. The communities of higher animals compared with 
SMOBE OF ANUS vc ok Fn PES i oo a 
Mutual warning of danger. Sentinels. Mutual “charity.” 
Nursing of the sick. Co-operation and division of labor. 
Common defense. ‘‘Fidelity and obedience” in animal so- 
cieties. Brief summary. 


CHAPTER II. 
WARFARE AND SLAVERY IN THE ANIMAL KINGDOM. 


1. Wars among higher animals...............ceceees 


Fables and stories of wars between troops of apes. The 


latter use no weapons or tools, 
viii 


iii 
a 
vi 


14 


20 


40 


Contents. 


PAGE. 


2. The war-like expeditions of the Amazon ant and the 


FOCmiea SOUQUINEE Gi A eins oh i a h 

Their tactics, the “humanity” of the victors. The military 
skill of the Amazon ants; their inability to feed themselves. 
The military tactics of the sanguine slavemaker (Formica 
sanguinea) discussed on psychological grounds. 


3. The pretended “automatism” in the psychic life of ants 

The individuals of one and the same colony act differently 

in combat. Acquired individual habits. Persecution of toler- 

ated or even of true guests. Ants tamable. Psychic influence 

of the numbers of a colony on their courage in combat. 
Marked ‘‘heroism” of individuals. Martial sports? 


4. The slave-making instincts of Formica sanguinea...... 
“Tradition and instruction” in insect communities. Experi- 
ments made with “self-taught” ants and bees. Seemingly 
intelligent plasticity of the instinct of slavery in Formica 
sanguinea, both with regard to the number and to the species 
of its slaves. On the nature of slavery in ants. The correct 
explanation is neither anthropomorphism nor mechanic automa- 
tism, but something between the two. 


* Other wats atid alliances of 'Shts oc cio. ccs dae ce oe vanes 
Fights between neighboring colonies. Alliances between 
hostile colonies, and their psychological explanation. Summary. 


CHAPTER III. 
ARCHITECTURE IN THE ANIMAL KINGDOM. 


1. A general survey of the architecture of animals........ 
2. The nests of ants. ve ota elek ealee age eae arse 
Various and wbincaee: forms of their nests. Nests of dif- 
ferent ant species. Psychological explanation of this difference. 
3. The nests of Formica sanguined........ cece ceeceeee ; 
Variety and plasticity in their forms. Variable panes of 
the nests of a colony. Periodic changes of nests; different 
places of residence in different seasons. These phenomena 
psychologically explained. Power of adapting the construc- 
tion of their nests to the surroundings, in order to secure 
them against hostile inroads. 
_ 4. How do ants build their nests?................. Reni. 
No rigid system of co-operation; seemingly intelligent self- 
determination. Suitable consideration of the different con- 
ditions of- temperature and moisture. Architecture of birds 
compared with that of ants; with that of mammals, in par- 


ticular of beavers. 
ix 


55 


67 


82 


103 


> 


Contents. 


PAGE. 


3. Other purposes, for which ants employ their architec- 
fibals: Ska ce ese RAN ere een Paes eect 


Stations, roads, galleries, stables for aphides, granaries, 
ramparts, etc. The grave-yards of ants in fiction and in 
reality. 

6. Is the building activity of ants guided by intelligence?.. 

Ants using their larvae as spinning-wheels. The “bridge 
building” of ants according to W. Marshall and Buechner. 
Experiments of Lubbock and Bethe. My own experiments. 
Results summed up. 


CHAPTER IV. 
BREEDING AND NURSING IN THE ANIMAL KINGDOM. 


1. A general view of the breeding instincts of animals ... 


Breeding and family life in animals with reference tc the 
preservation of species. This relationship psychologically 
explained. Breeding in social insects. The breeding instinct 
of ants dependent on the laws of organic development. 


2. Care of the young ‘among ants. 0.0.0.0 0s ce Ae 


Psychic characteristics of their breeding instinct and of its 
different manifestations. The differentiation of castes in 
bees and ants influenced by education. Intermediate forms 
between females and workers in ants, and their probable 
explanation. Pseudogynes and their relation to Lomechusa. 
Attachment of ants to their brood. Fond “mothers” and 
“aunts.” 


3. Adoption instincts in the animal kingdom...... canes 


Their frequent occurrence. Psychological explanation. Ants 
nursing Lomechusa-larvae. Different treatment of these 
larvae, both in normal sanguinea colonies and in such as 
contain pseudogynes. W. Marshall and the nursing of eggs 
of aphides by ants; “intelligent’? foresight of the future; 
comparison with the nursing of Lomechusa-larvae. Adoption 
instincts in birds and mammals. Results summed up; con- 
clusions, 


Conclusion. 30334 42.3. SD Gee TY & asa eo oe ee ares ; 


The automatism and the plasticity of instinct both in ants 
and higher animals reviewed. Man the only being in the 
visible creation gifted with reason. Humanization of animals 
is unscientific, and injurious to the moral order of human 
society, 

x 


153 


173 


192 


INTRODUCTION. 


VEN in ancient times, observers of animal life 

noticed that bodily size and psychic perfection 
are not always in direct proportion, but that the 
reverse is not unfrequently the case. Thus Aristotle’ 
declared that keenness of perception (rqv Tijs Siavoias 
dkpi(Beav) was often more manifest in smaller 
than in larger animals. Nor did it escape the great 
Stagirite, who was not only a logical thinker, but also 
a skilful observer, that many animals of low rank 
in the zoological scale were endowed, in some way, 
with a higher psychic life than the highest mammals, 
so much so, that its manifestations could be com- 
pared with human institutions only. He mentions, 
especially, ants and bees among those “bloodless” 
animals which possess a more intellectual soul than 
_ many animals of the other kind.?, The same thought 
was expressed by St. Augustine, one of the loftiest 
Christian minds, in the following terms: “We admire 
the works of the tiny ants and bees more than the 
bulky forms of whales.”’* And a distinguished modern 
naturalist, Emil Dubois-Reymond, has acknowledged 


1) “Hist. animal.,” 1. 9, c..7 (Becker I, 612). 

2) “De partib. animal.,” 1. 2, c. 4 (Becker I, 650). Aristotle’s 
division of animals into those with red blood and those with colorless 
blood in reality coincides with that of Vertebrates and Non-Vertebrates. | 
He uses the term “Bloodless Animals” for those which have no red 
blood. 

8) “De civ. Dei,” 1. 22, c. 24, n. 5 (Migne XLI, 792). 

1 


2 Introduction. 


in the name of his colleagues:t “With reverential awe 
does he (the naturalist) gaze at the microscopic speck 
of nervous substance, which harbors the soul of the 
ant with its industry, its instincts of architecture, 
order, fidelity and courage.” 

Surely, it was not without great reason that 
scientific observers of recent times applied themselves 
to the most careful and detailed examination of the 
life of ants, especially since the publication of Pierre 
Huber’s classical “Recherches sur les Moeurs des 
Fourmis indigénes’ (1810). Very many interesting 
facts of great value for psychological research have 
thus been furnished. However, dabblers in popular 
science, who viewed things from the standpoint of 
“vulgar psychology,” as Wundt termed it, misinter- 
preted these facts in a very unscientific manner; for 
they tried to draw conclusions from them which led 
to the humanization of animals, and denied the 
existence of any essential difference between the 
psychic faculties of man and brute. It is not so very 
long since Ludwig Buechner endeavored to pro- 
mote these ideas in his “Geistesleben der Tiere” 
(Berlin, 1876). As is generally the case with such 
shallow elaborations, Buechner has found not a few 
imitators and plagiarists. Therefore, it may not be 
out of place to examine these deductions from the 
standpoint of critical psychology. 

Sir John Lubbock, who, devoted himself to the 
study of ant life with the accuracy of a professional 
scientist, and who carefully refrained from the 


1) “Ueber die Grenzen des Naturerkennens.” Lectures by E. 
Dubois-Reymond, 1st issue (Leipzig, 1886), p. 127. 


Introduction. 3 


humanizing tendencies of modern times, states in the 
introduction to his book, “Ants, Wasps and Bees,” 
that ants rank next to man in the scale of intelligence, 
and that in psychic faculties they approach nearer to 
man than the Anthropoid apes even. George Romanes 
in the sixth edition of his book, “Animal Intelligence” 
(1895), devotes more than one hundred pages to ants, 
and thus indicates the great importance he ascribes 
to their psychic qualities. 

Prior to Lubbock’s work on ants, another prom- 
inent investigator of ant life, Dr. Augustus Forel, in 
his “Fourmis de li Suisse” (1874) had expressed 
the opinion that the principal factor in the psychic 
activity of ants was not individual intelligence, but 
social instincts (p. 444). Although he pretends to 
find even among ants remarkable proofs of intellect, 
he maintains that it cannot compare with the individual 
intelligence of the higher Vertebrates (as apes, seals, 
elephants, etc.). Most of my critics, likewise, espe- 
cially Forel and Smalian, in discussing my book 
“The Compound Nests and Mixed Colonies of Ants,” 
conceded that ants were guided in their life and doings 
almost exclusively by their social instincts. With the 
higher Vertebrates, however, intelligence is said to 
preponderate gradually over instinct. This is postu- 
lated by Darwin’s theory of evolution, which other- 
wise would be unable to explain the mental evolution 
of man from the animal kingdom. The only possible 
explanation according to this theory is to assume . 
that, to the individual mammal-intelligence of the 
hypothetical ancestors of man, there was added, 
through the development of community life, a higher 


4 “Introduction. 


degree of perfection in their social instincts, and that 
thereby the higher animal was gradually transformed 
into man. ; 

The tenability of this assumption will be discussed 
in the following chapters. It is understood, that in our 
comparative investigation, we shall be guided, not by 
the postulates of evolutionist theories, but by the prin- 
ciples of critical psychology, set forth at length in our 
former essay, “Instinct and Intelligence in the Animal 
Kingdom” (Herder, St. Louis, Mo., 1903). 

Lately there has been invented a theory on the 
psychic life of ants, which is dtametrically opposed 
to the popular attempts at humanization. Alb. Bethe 
has tried to set down ants and bees as mere “reflex 
machines,” devoid even of the simplest sensitive per- 
ception and cognition, whilst he considers the intelli- 
gence of higher animals to be beyond all doubt. Thus 
he hoped to succeed in destroying the parallelism 
established by us between the psychic faculties of ants 
and those of higher animals, from which we had 
drawn the conclusion: we do not need ant intelligence, 
therefore neither animal intelligence. Bethe’s work is 
of undoubted value on account of its attack on the still 
wide-spread popular views regarding ants as intelli- 
gent, human beings in miniature.? His theory has 


1) “Duerfen wir den Ameisen und Bienen psychische Qualitaeten 
zuschreiben?” Bonn, 1898. (“Archiv fuer die gesamte Physiologie,” 
LXX, 15-100.) 

?) In this regard the “‘Betrachtungen ueber die staatlich lebenden 
Immen,” published against Bethe’s essay by Charles Sajé in 
“Prometheus” (10 Jahrg., 1899, Nr. 486 and 487), go far beyond what 
is admissible. Similarly the essay by Kienitz-Gerloff, “Besitzen die 
Ameisen Intelligenz?” in ‘Naturwissenschaftl. Wochenschrift” (XIV, 
1899, n. 20 and 21), 


Introduction. 5 


already, on another occasion,’ been subjected by us to 
a thorough discussion. Besides, in a longer essay, 
especially intended for professional zoologists,? we have 
since then perfected our former argument, proving 
that ants are no more mere reflex machines than dogs 
and apes are intelligent beings. The theoretical side of 
Bethe’s psychological views was also noticed in the 
second edition of our essay “Instinct and Intelligence 
in the Animal Kingdom” (chapters 7 and 8).* In this 
work, therefore, we shall return to them but occa- 
sionally, to show the fatal results of attempting to 
vindicate the intelligence of higher animals by com- 
pletely denying psychic activities in ants. 

1) “A new ‘reflex theory of ant life’? (Biolog. Centralbl., XVIII, 
1898, n. 15, p. 577-588). a 

2) “Die psychischen Faehigkeiten der Ameisen.’”’ Stuttgart, 1899. 


(“Zoologica,” Heft 26) p. 134 and foll. with 3 plates. 
3) p. 144 and following. 








CHAPTER I. 
COMMUNITY LIFE IN THE ANIMAL KINGDOM. 


1. A General Survey of the Forms of Animal Com- 
munities. 


HE multiplicity of bodily shapes apparent in ani- 

mals is not more remarkable than the variety 
found in their biological relations. The majority of 
animals, lower as well as higher, live singly, and only 
temporarily join other individuals of the same ‘species 
for breeding purposes; no lasting psychic tie unites 
them with others of their species. Other animals live 
in pairs during the breeding season, and remain united 
until the young are old enough to shift for themselves ; 
this is the case with most birds and mammals. If the 
offspring continue to remain with their parents, fani- 
ilies develop into herds, embracing the members of 
different, allied families. Thus, v. g., wild oxen and 
horses, the chamois, antelopes and many apes are 
gregarious animals. While real gregariousness is 
based on family ties in a wider sense and is mostly 
permanent, other animals flock together to form troops 
or hordes to undertake journeys in common, e. g., our 
migratory birds in autumn, the Scandinavian lem- 
mings, etc. Insects, too, gather into similar temporary 
masses of individuals of the same or closely allied 
species, assuming the form of migrating swarms. 
Migrating locusts are known to everybody; but also 

7 


8 Chapter I. 


butterflies, dragon-flies and other insects have been 
observed to form similar swarms. 

But few animal species are so perfect in their social 
organization, that the members of the family construct 
their habitations, rear their offspring and provide for 
their food in common. These are what Aristotle calls 
Zéa rodtixa, animals leading a well regulated social 
life, comparable, in a way, to the social life of man. 
These animals are chiefly the so-called state-forming 
insects, the social wasps, bees, ants and termites. With 
the two latter social life is carried to the highest degree 
of perfection found in the whole animal kingdom. 
True, also among birds, the social weavers (Ploceus) 
construct habitations in common, inasmuch as they 
build their nests close together, and beavers unite in 
colonies to build their dams, when different pairs are 
interested in raising the water level at the same spot. 
But what is wanting in the associations of higher ani- 
mals is co-operation, including some suitable division 
of labor for the rearing and nourishing of their off- 
spring. The combination of all these elements of 
social life is found only among the social insects, and 
in a prominent degree among ants. 

Viewed from the standpoint of comparative psy- 
chology, social is preferable by far to single life. In this 
connection, of course, we mean a social life based on 
social instincts, on the laws of sensitive cognition, and 
not merely a union caused by the laws of vegetative 
life, as is the case with certain animal conglomerates, 
as sponges, corals, polyps and many species of Tuni- 
cates. The bond, which unites the different individuals 
of these species to a colony, is entirely material. They 


Community Life in the Animal Kingdom. 9 


live together from. immediate, vegetative necessity ; 
for they literally grow as branches from a common 
trunk. As it is an immediate vegetative necessity for 
plants to bring forth twigs, leaves and blossoms, so 
mere vegetative necessity forces a colony of Siphono- 
phores to separate into different loosely connected 
individuals, some serving the purpose of nutrition 
(nutrient polyps), others of propagation (sexual 
polyps), of perception (perception polyps), of loco- 
motion (swimming polyps), and of protection (pro- 
tective polyps). To apply to the members of such 
colonies the term “persons” (eating persons, swim- 
ming persons, etc.), as Haeckel and several other 
zoologists have done, is evidently out of place, because 
this term implies a psychic independence which these 
animals do not possess. It would be more justifiable 
to conceive the whole growth of .Siphonophores as 
one individual of imperfect unity, consisting of various 
members, which, on account of their different func- 
tions can more fitly be termed “organs” than “persons.” 

The similarity of social life in the colonies of 
polyps and of ants is very slight and superficial. The 
latter, in opposition to the former, consists of indi- 
viduals organically separated and independent in their 
psychic activities. The members of an ant colony are 
complete individuals united to each other, not by the 
laws of vegetative growth, but by tmstinctive sym- 
pathy. This kind of co-habitation must indeed be 
- regarded as a higher manifestation of psychic life 
unknown among solitary animals. 

It is true, with the state-forming insects also, the 
instinctive association of the individuals of a colony 


10 Chapter I. 


is based on an organic, i. e., vegetative fact, namely 
on the eeicd oe from one and the same ecb 
called a “queen.” 

Honey-bees have never more than one queen in the 
hive, ants may have several of them. The instinctive 
dependence of bees on their queen is not so great as 
was formerly believed. Moreover, in the bee-hive the 
queen has essentially no other function than that of 
laying eggs; for the rest, her attitude towards the 
social activities of the colony is entirely passive; even 
when the bees are swarming the old “sovereign” is 
generally hurried along by the crowd of her “faithful 
subjects ;” she does not lead the expedition, neither 
does she determine its direction.t However, a swarm 
of bees deprived of their queen will disperse, because 
they have no common center of attraction, no point 
of crystallization, so to say, around which to form a 
new colony. In bee-hives the instinctive bond uniting 
queen and workers is closer than among ants, because 
the odor emitted by the queen exercises a far more 
powerful attraction? on the workers than in the case 


1) Abbé J. J. Kieffer communicated the following observations: 
“An old queen must often be actually forced out of the hive by the bees 
already swarming; sometimes the bees are gone, the queen being left 
behind in the hive. In other cases I observed that the old queen had 
dropped to the grotind; in spite of this, the bees settled at quite a 
different place on some tree, and suffered themselves to be put in a 
new. hive which, however, they soon left again, because the queen was 
missing.” 

2) How powerful is this attraction, can be gathered from an obser- 
vation made by Fr. Spillmann, S. J., in June, 1896. On catching a 
cluster of swarming bees, a few hundred workers had remained in the 
catching apparatus and could not find their way to the new hive. Led 
by their sense of smell, however, they clustered around a queen that 
had been lying dead on the ground for eight days, although it belonged 
to a different hive, 


Community Life in the Animal Kingdom. 11 


of ants. Thus the queen of the bee-hive becomes in a 
higher degree the principle of union for the workers 
of her colony and for the regulated exercise of their 
instincts. For this very reason but one full-grown 
queen is tolerated in a bee-hive, while in ant nests 
several may be found. What follows may ultimately 
account for the fact. 

The workers among ants live much longer than 
among bees. According to my observations, our 
Formica species, as a rule, attain an age of two, some- 
times of three years, whilst the workers among bees 
die after a few weeks or months. For this reason a 
colony of ants can continue to exist for several years 
without a queen, and even produce males through 
parthenogenesis. This longer duration of life with 
workers among ants may perhaps explain, why they 
are less dependent on their queen than bees, and fairly 
accounts for the fact, that, with ants, queens are 
needed as the unitive principle of the colony in a far 
inferior degree. Hence in a community of ants the 
number of generating, impregnated females may be 
almost unlimited. In a populous nest of the hill-ant 
(Formica rufa) near Exaten (Holland), I once found 
more than sixty full-grown queens. A similar num- 
ber I met with in a nest of the small, red stinging 
ant (Myrmica scabrinodis). In fact, by far the 
greater part of European ant species have, as a rule, 
several queens in every colony of long standing. 
With foreign ants the case is pretty much the same. 

A community of bees, therefore, having only one 
queen, may aptly be compared with a monarchy. But 
on account of the great number of oviparous mem- 


12 Chapter I. 


bers and the consequent greater independence of 
instincts in the single worker, an ant colony bears the 
stamp rather of democratic, republican, even socialistic 
institutions. Viewed from the standpoint of compara- 
tive psychology, the community life of ants is more 
perfect than that of bees, on account of the greater 
psychic independence of each individual. It is this 
quality of individual independence that lends to ant- 
states, among all associations of animals, the greatest 
resemblance to the political societies of man based on 
individual intelligence and free will. This resemblance 
is of course never more than mere analogy; but it is the 
highest degree of analogy known to exist between the 
social institutions of man and of the brute. Nor is the 
term “state” applicable to the social organizations of 
ants or, in fact, to any animal community, in any other 
than a metaphorical’ meaning; yet it applies more 
perfectly to ant states than to any other family of 
insects, and to insect states rather than to those of any 
other animals. 
_ Another important reason, why with ant colonies 
the use of the term “state” is comparatively more 
appropriate than with the social organizations of other 
animals is, because colonies of ants are often not 
merely “enlarged families,’ but contain also members 
of entirely different species which are hospitably shel- 
tered in the colony. Thus a simple ant colony comes 
to be a compound animal society. The above-men- 
tioned strangers are partly ants belonging to other 
1) On this point vide A. Espinas, “Des sociétés animales” (2e éd.) 
p. 372. Also Karl E. v. Baer (in Stoelzle, ““K. E. v. Baer und seine 


Weltanschauung” [1896], p. 300); W. Wundt, “Vorlesungen ueher die 
Menschen- und Tierseele,” 2d ed., p. 451. 


Community Life in the Animal Kingdom. 13 


species, living in the colony as “auxiliaries” or 
“slaves”; partly they are members of altogether dif- 
ferent orders of insects, especially of certain beetles, 
as the genera Atemeles and Lomechusa, which are 
accorded a friendly reception by the ants, are licked 
and fed, their larvae being reared by the ants as if 
they were the latters’ own.? This is a special form 
of community life (symbiosis), found nowhere else 
throughout the animal kingdom. Symbiosis is only 
equal to real community life, when the members 
engage in mutual psychic intercourse. Between a 
hermit crab and a sea anemone that settles on the 
former’s back, between a small fish (Trachichthys tuni- 
catus) and a large sea nettle harboring it within the 
circle of its tentacles,* there is a mutual relation (mutu- 
alism) useful to both of them, without, however, 
approaching any psychic intercourse, although the one 
instinctively looks for the other. There is a similar 
relation between ants and many of their tolerated 
guests, whilst their relation to their slaves and to their 
genuine guests attains a higher degree of psycho- 
logical intercourse and becomes real community life. 
Moreover, parasites, hostile intruders and indifferently 


1) See Wasmann, “Die zusammengesetzten Nester und gemischten 
Kolonien der Ameisen,” part II. 

2) See the “Autobiography of a Lomechusa,’ in “Stimmen aus 
Maria Laach,’”’ LII .(1897), 69, where the literature of the subject is 
enumerated. The number of the regular nestmates of ants and 
termites is rather considerable. Our ‘‘Kritisches Verzeichnis der 
*myrmekophilen und termitophilen Arthropoden,” published in 1894, 
already contains 1,246 ant guests and 109 termite guests, having the 
most various biological relations to their hosts. Since then many new 
species from all quarters of the world have been discovered and 
described. 

3) See “Zool. Anzeiger,”? Vol, XI (1888), n. 278, p. 240. 


14 Chapter I. 


tolerated cohabitants are found in the society of many 
higher and lower animals. They are present like- 
wise in the nests of social wasps, hornets and bumble- 
bees; but genuine guests (Symphiles), which, in spite 
of their morphological difference, are treated by their 
hosts as enjoying equal rights, as members of the 
family, are met with only among ants and termites. 
That stray chamois or steinbocks should join a herd 
of goats, is evidently something quite different from 
the fact that ants keep aphides and scale-insects as 
their milk cows, and tend even their eggs; or that 
they feed from their own mouths certain species of 
beetles, which on being licked afford the ants a special 
pleasurable sensation, herein treating them the same 
as they do their own comrades and larvae. The 
mutual social relationship which is here seen to exist 
between the animals of different species, and which 
we term Symphily (ow-¢iia) is by far more per- 
fect. Although, as we shall show later on, it is inti- 
mately connected with the instinct of adoption which 
occurs also among higher animals, the relation exist- 
ing between ants on the one hand and their slaves 
and genuine guests on the other, is nevertheless a 
form of perfect Symbiosis unparalleled among the 
Vertebrates. 


2. The Social Basis of Ant States. 


As was already indicated, the ultimate foundation 
of ant states is organic. It is organic, not only be- 
cause it is due to the descent from a common ovip- 
arous female, but more especially because it is con- 
ditioned, in its essential outlines, by polymorphism, 


Community Life in the Animal Kingdom. 15 


in other words, by bodily difference in the individuals 
of a colony. Ant states are organically divided into 
fixed groups of different “castes,” possessing different 
corporal and psychic qualities. These castes take their 
origin from the peculiar organic development of ants; 
they depend on laws of vegative growth, not on the 
intelligence and free will of individuals, as do the 
classes of human society. By far the majority of 
members of ant colonies consist, of course, of wing- 
less neuters, which go by the name of “workers” or 
simply “ants.” These workers are a secondary form 
of the female, the ovaries being stunted, while brain 
and instincts are all the more highly developed.t 
With many ants, especially with the genera Pheidole, 
Pheidologeton, Eciton, Colobopsis, etc., the workers 
are again divided into two more or less strictly sep- 
arated castes differing in bodily structure, namely: 
workers proper and soldiers, the latter possessing a 
comparatively huge head and formidable jaws. The 
wingless workers and soldiers are entrusted with the 
colony’s social welfare; it is their duty to build the 
nest, to tend the young, to gather provisions and to 
defend the community against hostile invaders, whilst 
the winged males and females attend to the propaga- 
tion of the species. After having been fertilized, 
which is generally done in the air during their nuptial 
flight, the females lose their wings and become 
“queens,” either founding new colonies or being taken . 
back by workers into their old nest for oviposition. 
The basis, therefore, of the so-called political con- 


1) Hence they cannot be simply called “stunted females,’ no more 
than the workers among bees. 


16 Chapter I. 


stitution of ants’ is in fact organic; it consists in the 
descent from one fertile female, and in the differen- 
tiation of the descendants into castes differing in 
bodily and psychic qualities, as a result of the very 
same specific fertility. The social bond, however, 
which unites the members of an ant colony and sep- 
arates them from other colonies of the same species, 
is psychic and instinctive. It is the feeling of fellow- 
ship, the instinct of sociality, resulting from common 
descent; it is, moreover, the instinct of imitation 
which urges the workers of the same colony to act 
in concert. This unity and co-operation is effected 
‘by means of a certain sensile feeler language: by a 
touch of their feelers thousands of members of a 
colony immediately recognize one another as belong- 
ing to the same community and effectually discover 
the intruder; by taps of their antennae they exchange 
their feelings and perceptions and thus draw the 
attention of other workers of their colony to the same 
work. The same feeler language is also the means 
of communication of ants in mixed colonies with their 
auxiliaries of other species, and of genuine ant guests 
with their hosts. 

This distinction between members of their own 
colony and those of others is effected by very delicate 
organs of smell? situated in the antennae. Members 

1) We mean here in the first place the simple ant societies which 
embrace no members of different species. 

2) We have already proven in our work, “Die psychischen Faehig- 
keiten der Ameisen” (“‘Zoologica,” 26th issue, p. 10-16), that there is 
not merely question of a “chemical reflex” (as Bethe calls it), but of 
a real sensitive perception. On the other hand, Lubbock’s experiments 
(“On the senses, instincts and intelligence of animals’? [London, 1889], 
p. 233 and foll.) have shown that an arbitrarily chosen sign or pass- 


word is equally out of place, as is evident from the fact that an ant 
which has lost its feelers is nevertheless recognized by her nest mates. 


Community Life in the Animal Kingdom. 17 


of the same colony have the same delicate “nest 
odor,” and by licking strangers they are able to trans- 
fer it to other insects. A beetle of the genus Atemeles 
having been licked in a friendly manner by but one 
ant of a Formica colony, will be acknowledged as a 
friend by the other ants of the same colony, whilst 
otherwise they would attack it The “nest odor” can 
be communicated to members of other colonies not 
only by licking but also by feeding. The smell of the 
salivary gland secretions* thus seems to serve ants as 
well as bees as a means of recognizing the “citizens 
of the same state.” 

It is, no doubt, downright nonsense for Buechner* 
to put ant states on the same level with human 
republics, much more so to consider them more perfect 
than the latter. And when modern sociologists* try 
to establish their reforms of human society on such 
foundations, we are justified in styling their endeavors 
utopian schemes. The promoters of such ideas for- 


1) More on the significance of the salivary gland secretions as a 
means of recognition among ants will be found in the essay mentioned 
above “Die psych. Faehigkeiten der Ameisen,” p. 16 and 97 ff. On the 
latter pages we have also shown that it is not merely the smell of the 
salivary gland secretions adhering to a beetle, that induces the ants to 
receive it after it has been licked by a single ant of that colony, but 
that, besides, other psychic elements are in play and must be considered 
in explaining the fact. See also “Instinct and Intelligence in the 
Animal Kingdom,” p. 158. 

2) See the interesting little essay by N. Ludwig, ‘‘Futtersaft oder 
thierische Veranlagung als der Beherrscher und Ordner geheimniss- 
voller Vorgaenge im Bienenvolke,” published by the “Leipziger Bienen- 
_zeitung,”’ 1896. Likewise N. Ludwig, “Ueber Geruchempfindung und 
Riechorgan der Honigbiene” (‘“‘Natur und Offenbarung,” 1899, 9th 
issue, p. 554 ff.). 

3) “Geistesleben der Tiere,” p. 52. 

4) See e. g. Cognetto de Martiis, “Le forme primitive nella 
Mate vs economica.” Torino, 1881. 


18 Chapter I. 


get, that with man class differences rest on far dii- 
ferent bases than differences of castes among ants. 
With man they are the outcome of changeable, out- 
ward conditions of life, or perhaps the result of the 
intelligent free choice of the individuals concerned; 
with ants, however, they spring directly from the 
hereditary organic laws of polymorphism. Besides, 
those socialistic theorists forget that among ants there 
exists perfect equality and fraternity between all the 
members of a colony, for the very reason that these 
animals are guided by their social instincts only, not 
by independent reasoning, and that they therefore 
are never liable, as men unhappily often are, egotisti- 
cally to prefer their individual welfare to the common 
weal. If those socialist enthusiasts could transform 
men into ants, then they might be justified in pro- 
posing ant republics as the ideal political condition. 
H. E. Ziegler’ is right, therefore, in saying: 
“With ants the social differentiation is conditioned 
by organization and instincts, and is thus accurately 
fixed and regulated, whilst with man the social differ- 
entiation is due to education, exercise and custom; 
only the foundation of man’s social life is determined 
by certain social instincts, its further development, 
however, is regulated by the intellect, by education 
and custom . . . . To argue about man’s social 
institutions from the relations existing among insects 
would be committing a gross error, all the more so, 
if one should consider the communistic insect ‘states’ 
1) “Die Naturwissenschaft und die socialdemokratische Theorie,” 
p. 186. See also R. Leuckart, “Ueber den Polymorphismus der 


Individuen oder die Erscheinungen der Arbeitsteilung in der Natur,” 
Giessen, 1851. 


Community Life in the Animal Kingdom. 19 


as models of human communism.” Smalian' agrees 
with Ziegler on this point, and I hardly believe that 
any intelligent naturalist will dispute their position. 

But now let us examine the other conclusions these 
statements imply. The social life of ants in spite of 
its differing essentially from the human state, is 
nevertheless the highest degree of community life in 
the whole animal kingdom; even the social relations 
among the highest apes are far from reaching the 
perfection of ant states. The foundation indeed of 
social life and division of labor in ant states is organic, 
and is to a certain degree predetermined by nature 
with aprioristic necessity through bodily polymor- 
phism. Nevertheless, also with them the actuation 
of the social instincts is guided and determined in its 
details by the sensile cognition and experience of the 
individuals. Whoever falsely styled this individual 
sensuous experience of higher animals, such as dogs, 
apes, etc., intelligence,” should not be so inconsistent 
as to deny to ants a high degree of the same “individ- 
ual intelligence.’ Whoever without previous critical 
analysis of his psychological notions maintains down- 
right, that associations resulting from the sensile ex- 
periences of the individual are intelligent, must credit 
ants not only with the highest development of the 
social instincts, but also with the highest development 
of intélligence found in the animal kingdom. This 
we wish to prove more in detail. 


1) “Altes und Neues aus dem Leben der Ameisen,” in ‘“‘Zeit- 
schrift fuer Naturwissenschaft,” LXVII (Halle, 1894), 39. 

2) Which is done by Ziegler and nearly all modern zoologists, as 
we have shown in “Instinct and Intelligence,” chapt. 2. 


20 Chapter I. 


In what does the pretended psychological superi- 
ority of the associations of higher animals over ant 
states consist? Let us try to clear up this question. 


3. The Communities of the Higher Animals Com- 
pared with those of Ants. 


Both Ziegler and Darwin’ point to the fact, that 
the higher mammals, especially apes, “sometimes form 
societies for the purpose of receiving notice of danger, 
for providing mutual protection and defense, for ob- 
taining nourishment, sometimes even for united at- 
tacks on their prey.’* Societies of ants have the 
very same end in view. Although their main pur- 
pose is to rear their young in common, yet those 
other secondary purposes are not only not excluded, 
but their pursuit and attainment by ants reach a de- 
gree of perfection unequaled by the above mentioned 
higher animals. However, neither Darwin, nor Es- 
pinas, nor Ziegler, nor, in fact, any modern student 
of animal psychology has ever succeeded in proving 
that apes are conscious of their purpose, and therefore 
act with intelligence, and that ants are without con- 
sciousness of purpose, and therefore acting merely 
from instinct. 

Let us consider more closely the different points 
of comparison. The higher animals living in hordes 
aid their comrades by certain calls, giving warning 
of danger. Some of them, e. g., the chamois, post 
regular “sentinels” for this purpose. However, ants 
do the same and in a manner much more indicative 


1) “Descent of Man,” I. Chap. 4, 
2) Ziegler 1. c., p. 189. 


Community Life in the Animal Kingdom. 21 


of intelligence. The whole difference lies in the fact 
that instead of calls, the ants use another means of 
sensile communication, namely, their feelers. If a 
troop of “sanguine slavemakers,” as McCook calls 
them, (Formica sanguinea), approaches a nest of the 
negro ant (Formica fusca), then the first black ant 
which has noticed the foe hurries back into the nest, 
communicates her own fright to the other workers by 
rapidly tapping them with her feelers and thus gives 
a general alarm. The larvae and pupae are hurried 
down from the higher parts of the nest into the deeper 
galleries and chambers, and if the foe advances as far 
as these apartments, the black ants run head over 
heels through the secret openings at the opposite side, 
and with their precious burden climb up stalks and 
bushes to save it from the foe. Sometimes they re- 
sort to this final means of escape at the first news 
of danger and take to their heels before the van: 
guard of the foe has reached the interior of the nest. 
In a similar way, but adopting different tactics, the 
yellow and the brownish-black meadow ants (Lasius 
flavius and niger) struggle for safety, when their 
nests are attacked by some Formica species. As soon 
as the approach of the foe is discovered, the fact is 
announced with lightning-like rapidity throughout 
the colony by rapid strokes of the feelers. The 
larvae and pupae, the winged males and the queens 
are carried to the lowest recesses of the nest, and the 
‘avenues to it are hastily blocked up with earth to 
prevent the enemy’s advance. Whilst the small 
Lasius is constantly closing up the approaches to the 
interior of the nest with bits of earth, such of the foe 


22 Chapter I. 


7 


as have ventured too far, are seized and killed by 
_ crowds of the assailed. 

If with higher animals it is a mark of intelligence 
to “utilize the senses of all for the protection of the 
commonwealth,” the same must be said of ants, and 
in a more perfect degree. The posting of sentinels 
for the protection of the community may be observed 
with these social insects just as well, and even better 
than with the social apes. In a nest of Formica san- 
guinea comprising four species of slaves (or auxil- 
iaries), namely, F. fusca, rufibarbis, rufa and pra- 
tensis, which is under my observation for many years, 
I can verify this fact every day. We _ subjoin a 
diagram of this observation nest, as it will often be 
referred to in the sequel. 

The main nest and its annex are made of glass 
plates in wooden frames. The space between the two 
plates in each nest is partly filled with earth, their 
vertical distance being from Io to 12 millimeters, so 
that the ants have freedom of motion to perform their 
work without being able to screen themselves from 
observation. The upper glass plate is generally cov- 
ered with a black cloth; for if light were permitted 
to enter, the ants would coat the lower surface of the 
glass with earth in order to darken the interior of 
the nest. By means of glass tubes the main nest and 
its annex are put in communication with each other 
and with the other parts of the nest, which are like- 
wise of glass. (See diagram.) 

In the main nest, which corresponds to the interior 
of an ordinary ant nest, the majority of the ants are 
to be found with their queens, their larvae, pupae and 


23 


Community. Life in the Animal Kingdom. 


feeding tube. 


f 





aunex. 





— 

















main nest. / 





Fig. 1. 


24 Chapter I. 


guests. In the front nest we see generally a number of 
ants basking in the sun or engaging in different labors. 
In the top nest a small number of sanguinea, rufa and 
pratensis are usually found either on guard, or wait- 
ing for the flies or other food which I occasionally 
throw in. In the glass bulb of the feeding tube, even 
if it happens to contain no sugar or honey, there are 
always one or two ants, mostly fusca or rufi- 
barbis, which have a special liking for this depart- 
ment. Even on the dumping grounds, whither the 
ants carry their dead, there are, as a rule, a few ants 
to be found, remaining immovable and watching for 
any suspicious circumstance in the nest or in its 
vicinity. On March 26, 1896, from morning till eve- 
ning one F. fusca and one F. pratensis were posted in 
the refuse nest; on March 27, at 7 a. m. two F. pra- 
tensis; at 10 a. m. two F. sanguinea took their place. 
On March 28, in the forenoon, one sanguinea 
was on guard, which having been taken out by 
me and confined was soon after replaced by another 
sanguinea for the rest of the day. On March 29 the 
whole day one sanguinea; on March, 30 at 7:30a. m., 
two fusca; when at 8 a. m. I had taken out and con- 
fined one of the two, I found that within the space of 
half an hour another fusca had taken its place, where- 
upon both remained there during the whole of the 
forenoon, etc. Only during winter, after I had com- 
pletely emptied the refuse nest and left it in the 
same condition for a long time, no ants were seen 
there for several weeks, because this part of their 
nest was no longer $f any vital interesi to them, and 
because the cool temperature kept them in the main 


Community Life in the Animal Kingdom. 25 


nest and in its immediate vicinity. It can hardly be 
maintained that this posting of sentinels in the dif- 
ferent parts of the nest was merely due to poly- 
morphism; for the cognitive and appetitive powers 
of the single individual ants of those five species in 
various ways take a prominent part in it. As we 
shall show hereafter, the same obtains in other forms 
of the division of labor in ant states. 

“Social animals perform many little services for 
each other; horses nibble, and cows lick each other 
wherever they feel an itching; monkeys hunt for 
each other’s external parasites,” etc. Thus Ziegler 
reproduces the statements of Ch. Darwin. But ants 
of the same colony are quite as serviceable to each 
other. Whoever has kept ants in suitably arranged 
nests of observation, where they feel comfortable and 
at home, can observe such “acts of charity” a hundred 
times a day. Every time I gently lift the black cloth, 
which protects the upper glass plate of the main nest 
from the rays of the sun, I witness one or more of 
these lovely scenes. Just now a worker of F. san- 
guinea is lying immovable, stretched on her side, whilst 
some of the companions are washing her; a sanguinea, 
a fusca and a rufibarbis perform this work, and lick 
her carefully, whilst she continues immovable; then 
they turn her around and lick her just as carefully on 
the other side. After half a minute the light which 
floods the nest interrupts the performance, and they 
flee to some darker spot, the patient soon following 
their example. All the workers of each of the five 
ant species living in my mixed colony without distinc- 
tion render these services of cleanliness to one another. 


26 Chapter I. 


Sometimes one of the dominant, sometimes one of the 
enslaved species is the recipient, no distinction being 
made between masters and slaves in performing these 
offices. Just as with cows in licking each other, so with 
ants, the performance of this service generally causes 
no less satisfaction to the active than to the passive 
partner, and, when apes look for each other’s parasites, 
we must, in order to arrive at a correct psychological 
appreciation of such “kind offices,’ not overlook the 
fact that apes devour with great relish the parasites 
discovered in the fur of their comrade. 

As regards these mutual cleaning services, ants 
and the higher social animals are pretty much on a 
par. The only difference is, that with ants they occur 
much oftener than with the latter. In both they pro- 
ceed, in the first place, from the desire for cleanliness, 
which is no doubt of an instinctive nature.t In the 
second place, they are due to the instinctive, mutual 
attachment between the members of animal associa- 
‘tions. The fact that ants clean a dust-covered com- 
panion by carefully “brushing” her down with their 
mandibles and licking her with their tongues, when 
viewed from the point of comparative psychology, 
finds its explanation in the same psychic motives as 
when “apes, after having rushed through a thorny 
brake, will examine each other’s fur and extract every 
thorn or burr.” To lick off the dust is, by itself, not 
more agreeable for ants; than it is for monkeys to 
extract the thorns. 

With ants the mutual attachment of nest mates 


1) See Ballion, “De Vinstinct de la propreté chez les animaux,” 
2d ed., Bazas, 1895, 


Community Life in the Animal Kingdom. 27 


goes so far as to make them carefully tend their 
wounded and sick: companions, which is not the case 
with gregarious mammals. The above-mentioned 
observation nest of F. sanguinea, on which I bestowed 
special care and attention, allowed me to observe 
several times, so as to leave no doubt of the fact, that, 
without distinction of masters or slaves, sick com- 
panions, or such as were paralyzed by the formic acid 
of hostile ants, were carefully nursed and licked for 
whole days, until they recovered. Forel, too, considers 
it a general rule, that ants nutse their sick or maimed 
companions.! | | 

Ants as “sick nurses” seemed so strange to me, 
that I was unwilling to admit the fact, until I observed 
it myself. The first time was on March 16, 1895. I 
had replaced in the main part of the aforementioned 
nest a sanguinea which had been paralyzed in one of 
the narrow glass tubes by an ejection of formic acid, 
and was scarcely able to move in spite of her con- 
vulsive efforts. At first her companions, on approach- 
ing, appeared to take no notice of her distress. Yet, 
after a short time, they began to examine her with 
their feelers, and then carried her to another part of 
the nest where the greater number were assembled. 
In this place the sick ant was lying for the whole day, 
surrounded by a number of masters and slaves (fusca) 
which, mostly in groups, busied themselves about her. 
They licked her carefully, turned her over and licked 
her again, examined her with their feelers and licked 
her once more. This method of medical treatment was 
attended with complete success. The patient had fully 


1) See Lubbock, “Ants, Bees and Wasps,” Chap. V, p. 88 fi. 


28 Chapter I. 


recovered by the next day, whilst without nursing she 
would probably have perished, as is generally the case 
with ants paralyzed by poison. 

If, therefore, on account of this “nursing,” Lub- 
bock and Romanes ascribe to ants a certain degree of 
“care and tenderness” lavished on their sick and 
wounded companions, they are right in so far as those 
actions are due to instinctive impulses, and not to the 
conscious affections of rational beings. For comparing 
the associations of ants with those of higher animals 
it may, at any rate, be of particular interest to notice, 
that such acts occur also among ants, notwithstanding 
their highly choleric temperament. 

“Yet, social animals also render more important 
services to one another; thus wolves and some other 
beasts of prey hunt in packs and aid one another in 
attacking their victims. The Hamadryas baboons turn 
over stones to find insects, etc., and when they come 
to a large one, as many as can stand round, turn it 
over together and share the booty. Social animals 
mutually defend each other.” This quotation from 
Darwin’s “Descent of Man” cannot justify Ziegler 
any more than his former arguments in concluding, 
that the community life of wild cattle, baboons and 
other mammals is more closely related to the social 
organisms of man, than that of ants. On the contrary, 
the manifestations of social life recorded above occur 
with ants even in far greater perfection. 

Ants, too, hunt in company, especially the so-called 
sanguine slavemakers (F. sanguinea and rubicunda), 
the red Amazon ants (Polyergus rufescens, lucidus 
and breviceps), and all the species belonging to the 


Community Life in the Animal Kingdom. 29 


Dorylide genera Eciton and Anomma. These are the 
dreaded legionary ants and driver ants of tropical 
America and Africa. The military expeditions of 
F, sanguinea are generally undertaken in small divi- 
sions of from twenty to fifty workers, with the purpose 
not only of robbing the neuter pupae of the slave 
species (Ff. fusca and rufibarbis), but often also of 
pillaging the nests of smaller ants belonging to the 
genus Lasius, the larvae, pupae and winged individuals 
of which are carried off to be devoured. During the 
time of the nuptial flight of Lasius niger, many san- 
guinea colonies are hunting in the vicinity of their nest 
for the heavy Lasius females which drop to the 
ground. Then either singly or with united forces these 
robbers pull their victims into their strongholds, 
where they are mercilessly slaughtered. On the after- 
noon of August 24, 1888, I witnessed such a typical 
hunting expedition of several sanginea colonies near 
Exaten (Holland), on the outskirts of a fir planta- 
tion. The road passing the nests was covered far and 
wide with sanguineas rushing upon every Lasius 
female that dropped from the air, as upon a welcome 
booty. Within the space of an hour I counted more 
than one hundred females of Lasius niger that fell 
victims to the hunters. 

The individual initiative of ants is manifested or 
such occasions in the same degree as with the higher 
mammals; whilst concerted action and _ suitable 
co-operation reach even greater perfection than with 
the latter. At any time a troop of our common red- 
backed hill ants (F. rufa) may be seen on some forest 
path, with combined forces dragging to their home a 


‘30 Chapier I. 


large, heavy dung-beetle (Geotrupes typhoeus) ; or a 
number of them are hauling to the nest a large beam— 
to our eyes it is but a broken twig—which is more 
than fifty times the weight of any single ant! Some 
pull in front, others push from behind, and even if the 
latter for a few seconds pull in the wrong direction, 
they soon notice it, and off it goes in the right direction 
to the nest. On April 25, 1897, I observed in the 
neighborhood of a pratensis nest near Exaten, two 
- workers dragging together a beetle of the genus 
Calathus towards their hill; they went at a double- 
quick, without hindrance on either side, both ants run- 
ning backwards with equal speed. 
The mode of acting in concert is different with 


different species of ants. Among our Formica species. 


it reaches its highest degree of development with the 
hill ants (F. rufa and pratensis), the initiative of the 
single ants bearing rather a secondary part. The 
sanguine slavemaker (F. sanguinea), however, which 
is able to proceed unitis viribus wherever it seems suit- 
able, combines with this power a remarkable degree 
of individual initiative, similar to that noticed in dogs, 
apes, and other higher animals. 

It is of special interest to watch the co- -opetation 


and division of labor of different species in mixed | 


colonies of ants. In my above mentioned observation- 
nest, which contains besides F. sanguinea four other 
Formica species as auxiliaries, these five species have 
divided the work necessary for the welfare of the 
community, so as to give each species exactly the share 
corresponding most to its instinctive preferences. This 
division of labor, however, is neither mechanically 


+ 


Community Life in the Animal Kingdom. 31 


defined, nor confined within the cast-iron rules estab- 
lished by the specific character of each ant, but the 
workers of one species will at least to some degree 
take part in the work of any other species. Thus e. g. 
the rearing of the young in the main nest (see p. 23) 
is chiefly attended to by the sanguineas themselves, 
but all the four auxiliary species join in the same 
work. In the glass bulb of the feeding tube containing 
the sugar, the greater number of visitors consists. of 
fusca or rufibarbis filling their crops by licking up 
sugar or honey, with which they hurry to feed their 
companions in the other parts of the nest. Sanguinea, 
rufa and pratensis often prefer to carry the grains of 
sugar “in the lump” from the feeding tubes to the top 
nest, manifesting again various individual differences 
in their mode of action, quite independent of their 
specific character. Sometimes the lump of sugar is 
carried only as far as y (see diagram on p. 23) to the 
top of the tube, whence it is transported by other ants ; 
generally, however, it is immediately brought (beyond 
x) to the front nest, where it is carried to the bottom 
in their mandibles, or else, but rather seldom, it is 
simply dropped from a considerable height. This i 
observed quite often with sanguinea, more rarely with 
pratensis. When I introduce a large fly or some other 
live victim into the front nest, it is mostly sanguinea 
and rufibarbis that dart upon it furiously, whilst rufa 
and pratensis manifest on such occasions remarkable 
‘ skill and perseverance in holding down the struggling 
and fluttering victim. The sanguinea with their 
powerful mandibles attend chiefly to the dismembering 
of their prey, whilst the conveyance of the larger pieces 


32 Chapter I. 


into the inner parts of the nest is generally done by 
rufa or sanguinea. 

It was of special interest to watch the behavior of 
my ants, when a new individual of the beetle Lom- 
echusa strumosa which lives with F. sanguinea as a 
genuine guest, was introduced into the top nest. At 
first, as long as there were no rufa and pratensis in the 
nest as auxiliaries, he was, if not descending into the 
main nest himself, taken at last by a sanguinea and 
carried down, in spite of his obstinate, passive resist- 
ance. Later on it was generally rufa and pratensis 
that transported the guest who attracted their attention 
in a remarkable degree. Once a rufa happened to be 
alone in the top nest and for a long time was trying 
in vain to get hold of one of the two Lomechusas 
happening to be there, when all at once she ran down 
to the front nest. Scarcely more than three seconds 
had elapsed before she returned with four other rufas 
which she had called to her assistance. Now the five 
rufas immediately set to work with united efforts 
to raise the Lomechusas, each of which was then car- 
ried down to the main nest by one of the ants.* 

If similar scenes had been witnessed in a society 
formed of different species of higher animals, we could 
not help admiring the harmonious co-operation and the 
suitable, but by no means mechanical, division of labor. 
However, it is not the higher animals, but ants that 
act in this way, and in order to save the pretended 
intelligence of the former, ants are classed as in- 

1) A more accurate description of the last-mentioned observation 
will be found in our essay, “Die psychischen Faehigkeiten der 


Ameisen” (‘‘Zoologica,”’. 26th issue, Stuttgart, 1899), p. 63 ff. in the 
chapter on the power of communication in ants, 


Community Life in the Animal Kingdom. 33 


stinctive automatons, or even as unperceptive “reflex 
machines” ! 

Yet, neither in ants nor in any other animals, are 
co-operation and division of labor such as to become 
mutual, individual assistance, as is the case with man. 
The same object attracts the attention of several indi- 
viduals and leads them to busy themselves about it, | 
each in its own way. Working in company is due 
partly to the similarity of instinctive dispositions in the 
single ants, partly to the instinct of imitation. H. v. 
Ihering refers to this in the case of the Brazilian leaf- 
cutting ants (Atta), and has pointed out the psycho- 
logical importance of this difference as it exists 
between societies of animals and man.* 

Everybody knows that not only the higher mam- 
mals but also the social insects unite in defending their 
community and especially their young. To be thor- 
oughly convinced of this fact you need but step on 
a wasps’ nest or sit down on an ant hill. Indeed, the 
perfect unity and heroic “self-sacrifice” which social 
insects and in particular most ants display in defend- 
ing their nests and their offspring are simply unsur- 
passed by any other animal. This “unselfishness,” 
this “spirit of sacrifice’ and “motherly love” in ani- 
mals will be referred to in particular, when we come 
to speak of the breeding and nursing instincts. 

-Higher gregarious animals, e. g., bisons or baboons, 
do not in defending the community against a common 
‘foe defend also the individuals as such. When a 
hunter lying in ambush has killed one of the herd, the 


1) “Die Ameisen von Rio Grande do Sul,” in ‘Berliner Entomo- 
logische Zeitschrift,” 1894, 3d issue, p. 346. 
3 


34 Chapter I. 


other bisons generally take an inquisitive sniff at the 
corpse, but they make no assault on the foe for the 
sake of a wounded or dead companion. Wolves are 
far more unceremonious. Instead of devising plans 
for revenge, they devour their dead or wounded 
“brother.” Ants, when engaged in common defense, 
aim at defending the individual of the colony just as 
little as do the higher animals. An assailed ant is 
never defended by her companions for her own sake. 
They rush upon the foe, only because they see in him 
a common danger, and because their warlike spirit has 
been aroused. This was noticed by Forel and Lub- 
bock, and I can only confirm it. Therefore, neither 
higher animals, nor ants, when either at work or in 
battle, manifest anything like individual assistance in 
the human sense of the word.* 

“All animals living in a body, which defend them- 
selves or attack their enemies in concert, must indeed 
be in some degree faithful to one another ; and those 
that follow a leader must be in some degree obedient. 
When the baboons in Abyssinia plunder a garden, they 
silently follow their leader ; and if an imprudent young 
animal makes a noise, he receives a slap from the 
others to teach him silence and obedience.” 

Examples perfectly similar to the one just men- 





1) The pretended instances of individual assistance in the legionary 
ant Eciton hamatum recorded by Belt (The Naturalist in Nicaragua, 
2d ed., 1888, p. 26), are easily explained by the fact that these migrat- 
ing ants tried to take along their straggling companions, as is often 
the case in migrations of European Formica species. Therefore, there 
is no reason why we should credit the Ecitons with a higher “‘sympathy 
for their companions” than other ants, as Romanes does = (‘Animal 
Intelligence,” 6th ed., p. 48). This case is no proof of sympathy, but 
merely a manifestation of the instinct of sociableness. 


Community Life in the Animal Kingdom. 85 


tioned, and which Ziegler has borrowed from Darwin’s 
“Descent of Man,” may, when purged of arbitrary 
anthropomorphic interpretations, be recorded also of 
ants. He who takes the terms “fidelity” and “obedi- 
ence” as they are applied to human beings, namely as 
reasonable, voluntary subjection to the demands of 
duty and authority, can ascribe “fidelity” and “obedi- 
ence” to baboons as little as to ants. From the point 
of view of critical psychology it is ridiculous to inter- 
pret the “slap” given to the young baboon by its senior 
to be, as among men, an admonition to fidelity and 
obedience. The imprudent cry uttered by the young 
baboon, if the story is to be credited at all, excited the 
instinctive anger of the old apes as they were silently 
advancing. The instinctive association of certain sen- 
sile perceptions with certain sensile impulses affords 
a much simpler and more natural explanation of this 
fact. If, therefore, Darwin and Ziegler on this account 
ascribe to baboons fidelity and obedience in the human 
sense,’ they are but arbitrarily humanizing the brute, 
as indeed Darwin has done time and again in the book 
quoted above. 

A slight analogy of what we call fidelity and obedi- 
ence may indeed be observed in many animals, not 
only in the higher species, but also in state forming 
insects. Wherever a certain individual is the center 
of operation for the instincts of the rest of the com- 
munity, the latter will show it fidelity and obedience. 
‘The swarming bees cluster around their queen 
“faithful and obedient.” This allegiance is, of 


1) See my former essay, “Instinct and Intelligence in the Animal 
Kingdom.” (Herder, St. Louis, Mo.) 


36 Chapter 1. 


course, not so prominent with ants, whose queen 
is much less of a center for the instinctive activi- 
ties of the workers. With ants it is just the 
workers that by their restless activity and the remark- 
able display of individual initiative, are most power- 
fully stimulating the instincts of their companions to 
imitation, and thereby to actual co-operation in a given 
work. The only difference between the baboons 
described by Darwin and our ants is, that with the 
former the instinctive communication between the 
single individuals of a troop is effected mostly through 
calls, with the latter, however, through taps of the 
feelers. But both sometimes resort to more drastic 
gestures to supplement their means of “communica- 
tion.” If an excited F. sanguinea or fusca can not 
‘succeed by taps of her feelers in inducing a companion 
to join her work, she sometimes seizes her by the man- 
dibles or by a leg and simply drags her to the object 
which had first attracted her own attention. In the 
same way an ant often protects her comrades from 
a threatening danger first noticed by her. In my 
observation nests I repeatedly noticed some F. san- 
guinea or fusca, by taps of her feelefs or some other 
more drastic measures warning their companions to_ 
be “on their guard.”” When, e. g., I took away the glass 
tube connecting the feeding bulb with the top nest 
(see diagram p. 23), and caught a few of the “senti- 
nels” that instantly sallied forth from the opening of 
the top nest ready to fight, I often remarked some ants, 
that were posted near the opening of the top nest, 
approaching the others, tapping them with their feelers 
as a danger signal, and even getting hold of one, that 


Community Life in the Animal Kingdom. 37 


was about to run out, and pulling her back from the 
dangerous spot. To interpret such psychic manifesta- 
tions in higher animals as “intelligent actions” is evi- 
dently inconsistent with denying to ants an equal or 
even higher degree of “individual intelligence.” Criti- 
cal psychology will regard such occurrences in ants 
as well as in higher animals merely as associations of 
sensile representations and impulses, which must be 
classed as instinctive sensation, and not as intelligent 
thought". The social instincts of animals, which in 
their actual use are variously influenced and ruled by 
individual sense experience, perfectly explain all the 
appearances of “fidelity,” “obedience,” “caution,” etc., 
which occur with state-forming insects not in a lower, 
but rather in a higher degree than with apes and other 
mammals. To credit higher animals with quasi-human 
intelligence is, therefore, to humanize animals in a 
manner equally arbitrary and inconsistent. 

To sum up the results of our comparative study on 
the social life of ants and of higher animals. The 
associations of apes and of higher Vertebrates are 
based on social instincts, which lead them to co-operate 
for mutual protection and defense, and partly, too, 
for the procuring of food. This co-operation is more 
or less powerfully influenced and varied in its manifes- 
tations according to the sensile experiences and affec- 
tions of different individuals. Exactly the same mode 
of co-operation, but of a still more perfect, suitable, 
‘and variable nature, we observe also in ant states. 
With these animals, too, it is founded on _ social 


99 66 


1) See “Instinct and Intelligence in the Animal Kingdom’ 
(Herder, St. Louis, Mo.), especially Chap, III. 


38 Chapter I. 


instincts, which, corresponding to their organic poly- 
morphism, are different in different classes (castes) 
of the state. As regards the application of the sensile 
experiences and affections of individuals, there exists 
within the range of these classes a very great inde- 
pendence and variableness of individual action, which 
with several ant species, e. g., the sanguine slavemaker 
(F. sanguinea), is scarcely inferior to that observed 
amongst higher Vertebrates. Besides, the perfection 
of social co-operation of higher mammals is far from 
equaling that of ants; for with the latter it extends 
not only to protection, defense and hunting, but also 
to construction of their dwellings, to the rearing of the 
young, and to the support of all the “members of the 
state” by comparatively few individuals, going by turns 
in quest of food and supplying the community with 
provisions. Nothing of the kind is known of apes or 
other higher animals. The providing of food_in par- 
ticular varies greatly with the different kinds of ants: 
it embraces “cattle herding” (the keeping of aphides), 
hunting (robbing of insects, in particular, robbing of 
the pupae of other ants), agriculture (grain gathering 
ants), horticulture (ants raising fungi), etc. Nor are 
the military expeditions of several ant species under- 
taken merely from want of food, but also for the sake 
of making slaves, the ravished pupae of workers of 
other ant species being reared as members of their 
own state. Through this suitable incorporation of 
outsiders into their own colony the community life of 
ants in the “mixed colonies” reaches a quasi-intelligent 
universality, which is vainly sought for among higher 
animals, The same universality is manifested also by 


Community Life in the Animal Kingdom. 39 


the fact, that many ant species treat like members of 
their own family even different orders of insects, 
namely the beetles of the genera Atemeles, Lomechusa, 
Xenodusa, etc., which are known as “genuine ant 
guests,” and that they even tend and rear their young 
as if they were their own. 

It must, therefore, be conceded that the community 
life among ants is more developed and: more perfect 
than that among apes and other higher animals ; hence, 
from the point of view of comparative psychology, 
the communities of ants represent the most perfect of 
animal societies. 


CHAPTER II. 
WARS AND SLAVERY IN THE ANIMAL KINGDOM. 
1. Wars Among Higher Animals. 


ODERN evolutionists represent the social life 
among higher Vertebrates with the aim of mak- 
ing it the main support of the bridge spanning the 
chasm between man and the brute. Thus, Ziegler 
concludes his description by a psychological parallel, 
in which, just as Darwin did, he tries to establish the 
greatest possible similarity between the social life of 
animals and of man. Let us examine this evolution- 

istic attempt in the light of scientific psychology. 
Says Ziegler : “There exists, therefore, among ani- 
mals a social community life similar to what we meet 
among the hordes and tribes of uncivilized nations. 
Even wars, which have taken place among the hordes 
and tribes of the human race since prehistoric times, 
have their counterparts in the animal world, as is 
proved by the following example recorded by Darwin. 
“Brehm states on authority of the well-known 
traveler Schimper, that in Abyssinia, when the baboons 
belonging to one species descend in troops from the 
mountains to plunder the fields, they sometimes 
encounter troops of another species and then a fight 
ensues ; the geladas roll down great stones, which the 
hamadryas try to avoid and then both species, making 

a great uproar, rush furiously against each other.’ ” 
How far the “therefore,” which should connect 

40 


/ 


Wars and Slavery in the Animal Kingdom. . 41 


the social life of animals and of man, is supported by 
facts, has been shown in the preceding chapter. Even 
in higher mammals individuals of social communities 
co-operate merely as far as their social instincts guided 
by individual sensile experience will allow. In man, 
however, community life is due to social instincts 
as to its foundation only, but in its perfect develop- 
ment to the intelligent, free self-determination of indi- 
viduals. Ziegler and Darwin are far from having 
furnished the proof, that the latter element occurs also 
in higher animals. Or do they perhaps think that 
the wars which hordes of apes wage against each other 
contain this proof? Let us see. 

In the above description it is stated that the baboons 
roll down stones-at their enemies intentionally, and 
thus, as it were, use the stones as weapons, as e. g., 
in 1809 the Tyrolese occasionally did in their struggle 
for liberty against the French and Bavarians. But 
regarding apes, the statement is a myth. Pechuel- 
Loesche corrected the passage in the third edition of 
Brehm’s “Tierleben.” “We are told,” he writes, “that 
apes defend themselves with broken branches, and it 
is pretty generally assumed that they hurl down on 
their opponents stones, fruits, pieces of wood and other 
objects. This belief is probably due altogether to 
inaccurate observation... Its originators and abettors 
have perhaps seen only, what they from various 

1) We sincerely regret that Mr. Pechuel-Loesche was not allowed 
to subject the 3d ed. of Brehm’s ‘“‘Tierleben,’”? which he revised, to a 
thorough psychological revision. Although several of the most offensive 
passages were corrected or omitted, yet Brehm’s peculiar style has not 
changed; he cannot possibly refrain from intentionally humanizing the 


brute. See a criticism of this work in “Natur und: Offenbarung,” 
XXXVII, 570. 


42 ‘Chapter II. 


accounts supposed to be the fact, not what took place 
in reality. Apes living in trees, in wanton playfulness, 
break off withered branches by jumping on them, by 
snapping and shaking them ; but they do not throw 
them at a person who stands below. Neither do they 
throw fruits or other objects which they hold in their 
hands; they rather drop them quite naturally on being 
-frightened or put to flight. Moreover, baboons, among 
which I was able to observe especially the tschakmas, 
often watching hundreds of them very carefully, never 
think of throwing down stones from their rocky eleva- 
tions at their pursuers. It is true, from the place where 
they happen to be, stones sometimes roll or fall down, 
but merely by chance and also at times, when no enemy 
is in sight. . . . Together with my wife, who 
derived great pleasure from watching the behavior of 
the baboons,—they were often the only living beings, 
and very noisy at that, in the rocky deserts of South- 
western Africa,—I have minutely studied their doings 
precisely on this head to convince myself whether they 
actually throw. They assuredly do not.” 

What light is thrown by these critical observations 
of Pechuel-Loesche on the “individual intelligence” 
of apes so highly prized by modern evolution? Light 
enough, indeed, but extremely compromising for that 
theory. In spite of their highly developed brain, which 
in anatomical structure bears the closest resemblance 
to the human brain, apes are nevertheless unable to 
draw even the simplest conclusions, which might lead 
them to the use of branches and stones as weapons. 
The spider weaving its ingenious web to ensnare its 
prey, or casting out silky threads to entangle its vic- 


Wars and Slavery in the Animal Kingdom. 43 


tim; or the “ant-lion’” (Myrmeleon formicarius), an 
insect belonging to the Neuroptera, the larva of which 
from the center of its funnel-shaped sand-pit with its 
long and flexible mandibles hurls particles of sand at 
an ant, to bring her to the bottom of the pit ; or the 
archer fish (To.rotes iaculator), directing jets of water 
upon small insects resting on aquatic plants, thereby 
bringing them down into the water as his prey : these 
animals, so low in the zoological scale, are far nearer 
to man, as regards the suitable use of weapons, than 
the highest apes, although on evolutionistic principles 
the latter only could possibly form the transition 
between man and the brute. That apes through their 
imitative instincts and by human training are able 
to “learn” the use of a few simple tools, only proves 
the intelligence of man and the power of sensitive per- 
ception in apes. Had apes themselves but a trace of 
intelligence, they would have invented long ago even 
in their free state of nature the use of a few simple 
means of defense, such as branches and stones. But 
why did they not? The only possible, scientific answer 
is : because they evidently have no intelligence. Not 
the brain alone makes man an intelligent being, but 
his spiritual soul, and this spiritual soul is wanting in 
the highest apes as well as in insects. True, modern 
evolution is fond of ignoring all facts, which will not 
fit into its tissue of hypotheses. But we need not say 
that such a proceeding is highly unscientific. 
Darwin’s and Ziegler’s attempt at putting the wars 
of apes on a level with those of uncivilized tribes, has 


1) See “Instinct and Intelligence,” etc. (Herder, St. Louis, Mo.), 
p. 60 and 160. 


44 Chapter II. 


proved unsuccessful. Even the most savage nations 
employ tools and weapons of various kinds in order 
to catch their prey or to wage war against their foes. 
The parallel drawn by Darwin and Ziegler between 
the wars of apes and of savages proves to an unpreju- 
diced observer the very reverse of what Darwin and 
Ziegler intended to prove : it proves the essential dif- 
ference between the merely sensitive, psychic faculties 
of the highest vertebrates, and the spiritual, mental 
faculties of man. 


2. The M ilitary Expeditions of the Amazon Ant and 
of the Sanguine Slavemaker. 


The wars of ants bear far greater resemblance to 
human wars than those of the apes. Indeed, ants no 
more than other animals use any other weapons than 
those furnished by nature, namely their swordlike 
mandibles, their poison stings and poison syringes, 
but they use them in a manner which of all animal 
combats most resembles human strategy. Whoever 
watched a military expedition of the red Amazon ants 
(Polyergus rufescens) or of the sanguine slavemakers' 
will no longer entertain any doubts on the subject. 
The Amazon ants, the European Polyergus rufescens 
as well as the North American P. lucidus,? advance 


1) Since the issue of the book, “Die zusammengesetzten Nester 
und gemischten Kolonien der Ameisen” (1891), I have had occasion in 
Lainz near Vienna to observe a number of other Polyergus expeditions, 
and besides, several sanguinea expeditions near Vienna and in Lim- 
burg (Holland), etc. 

2) Called by McCook the “shining slavemaker,’”? whose habits he 
observed near the Allegheny mountains. There are still three other 
subspecies (races) of P. rufescens found in N. America, P. breviceps 
Em., bicolor Wasm. and mexicanus For. 


Wars and Slavery in the Animal Kingdom. 45 


on the war-path in large serried columns, the sanguine 
slavemakers, however, the European as well as the 
North American,’ in smaller, less serried detachments ; 
both, but especially the Amazons, try to storm the 
hostile nest by a fierce attack, and to stun the numeri- 
cally superior foe and to put him to flight by the 
suddenness of the onslaught. Great success generally 
attends these tactics. Forel, in his ‘““Fourmis de la 
Suisse” (1874), p. 306, has several similar instances, 
some of which we wish to bring to the notice of the 
reader. When Forel brought a bag containing a 
whole colony of meadow ants (F. pratensis), which 
in size and strength surpass the Amazons, into the 
neighborhood of an Amazon nest, several of the 
Amazons at first dashed fiercely into the midst of their 
numberless enemies; twenty of them were as a rule 
sufficient to rout fifty times that number of pratensis. 
Another time an army of Amazons just returning 
from the pillage of a slave nest were depositing their 
spoils of ant pupae in their nest; previous to setting 
out on a new expedition, when Forel at a distance of 
one meter from their nest and in the path of their 
expedition emptied a large bag of F. pratensis. In 
three minutes the whole army of the Amazons had 
encircled the hostile camp appearing quite unex- 
pectedly. They stormed it in an-instant, drove out 
the pratensis and ransacked the nest for its cocoons.—I 
would like to hear of apes ever displaying similar 
‘military skill. 

- It is characteristic of the military tactics of those 


1) Formica rubicunda and integra Em, are the principal N. Ameri- 
can races of the European Formica sanguinea. 


46 Chapter II. 


ant species which undertake slave hunting expeditions, 
to kill the hostile ants only when resistance is offered. 
Fleeing F. fusca or rufibarbis are pursued merely to 
obtain the larvae and pupae which they are carrying 
off; booty, not slaughter is the object of the victors. 
If apes or other higher animals were to act similarly 
in their wars, then our modern advocates of evolution 
would not fail to make the following reflections: 
“Here we find the first traces of genuine humanity, 
which shrinks from unnecessary bloodshed; what 
these animals consciously aim at is, not to fight, but 
to gather the fruits of victory,” etc. In ants such 
reflections are readily granted to be ridiculous human- 
izations of the brute; but never would it be conceded 
in the case of apes, not because the psychic manifesta- 
tions are really different, but rather to safeguard the 
evolutionistic theories. 

The military skill of the Amazons (Polyergus) is 
no doubt unexcelled amongst ants, but also amongst 
other animals. It is even far superior to the military 
tactics of the sanguine slavemaking ants, although the 
latter manifest in their whole character a more perfect 
development of what is called “individual intelligence,” 
i. e., the suitable application of their sensitive experi- 
ences. But the Amazons in private life are the dullest 
and most awkward “instinct beings’ you can imagine. 
Although they are able to take liquid food by licking 
just as other ants, they have nevertheless almost totally 
lost the instinct of feeding themselves, and would 
starve, unless they be fed from the mouth of their 
slaves. This fact makes it quite evident, that even in. 
the grandest military exploits of the Amazons there 


Wars and Slavery in the Animal Kingdom. 47 


enters not the slightest trace of genuine intelligence, 
but only instinctive sensitive faculties; for, an animal, 
that even in a state of utmost destitution is unable to 
combine his feeling of hunger with the perception of 
nourishment and the impulse to eat, can surely not 
be credited with even the lowest degree of delibera- 
tion. “A being that is physically able to eat, but has 
lost the habit of tt, is the greatest libel on animal intel- 
ligence.’’* 

Against this conclusion Dr. Smalian? has raised an 
objection which we are now going to examine. He 
believes our argumentation unsound; and asks “How | 
does Wasmann know that the Polyergus are at all able 
to feed? The basis of his argument is in concluding 
from the nature of the eating organs the ability to 
eat. And he states, that he has once seen Polyergus 
taking food independently; however, the matter is 
doubtful; for in the case of animals which otherwise 
never feed themselves but are always fed by others, 
it is impossible to know, whether the food they 
touched was actually consumed.” 

That Smalian should make such an objection may 
be explained only by assuming that he does not know 
the mode of life of Polyergus from actual observa- 
tion; otherwise he would hardly have been led to attack 
our argumentation. Besides, he has not reproduced 
in full the proofs which he controverts. Indeed, it 
was also from the anatomical structure of the mouth- 
parts of this ant that we drew the conclusion, that no 
organic impossibility prevented the independent feed- 

1) “Die zusammengesetzten Nester und gemischten Kolonien der 


Ameisen,”’ p. 204. : 
2) “Altes und Neues aus dem Leben der Ameisen,” p. 42, 


48 Chapter II. 


ing of Polyergus. Our chief argument, however, was 
the biological fact, that the Amazons do really some- 
times lap up liquid food, if by chance it comes in con- 
tact with the lower parts of their mouth. Dr. Smal- 
ian has undervalued this fact. Not only once, but 
repeatedly I saw and followed it up with a lens, how 
some Amazon which had pierced an ant pupa with its 
mandibles, licked up with her tongue the fluid flowing 
from the wound, and sometimes spent a considerable 
time in this occupation. Now, since the reception of 
food in ants generally takes place by licking, it is hard 
to understand, why it should be impossible to know 
in this case, whether the food has “actually been con- 
sumed.” 

-Besides Dr. Smalian has failed to notice Adlerz’ 
observations mentioned in the very passage quoted by 
him. Like myself, Adlerz has witnessed that the 
Amazons frequently lick up the moisture condensed on 
the glass walls of their artificial nests.. That Amazons 
are able to feed independently is, therefore, an estab- 
lished fact which can not be done away with. Why, 
therefore, do they starve, when they are confined in 
a test tube together with some honey or some appetiz- 
ing ant pupae, but separated from their slaves by which 
they are wont to be fed? The only possible and psy- 
chologically correct answer is : because their hunger 
does not compel them, like other animals, to seek for 
food themselves, but only to beg food of other ants 
by taps of their feelers. The sensitive perception of 
the food placed immediately before them, in spite of 
their feeling of hunger does no longer excite in them 
the natural imffulse of tasting it. With these ants 


Wars and Slavery in the Animal Kingdom. 49 


the instinct of independent quest of food and of its 
independent reception has perfectly degenerated. They 
have become utterly dependent on their slaves. Once 
more we ask Dr. Smalian and other friends of animal 
intelligence: Is it possible that a being, which pos- 
sesses but a trace of intelligence, should no longer be 
able to combine the sensitive perception of its proper 
food with the feeling of hunger? Therefore we still 
maintain: A being that ts physically able to eat, but 
has “unlearned” the habit of it, is the greatest libel 
on animal intelligence. 

The brilliant military talent of the Amazons is, 
therefore, a merely instinctive power, which is assisted 
by no individual intelligence. Just the most wonderful 
manifestations of the psychic life of animals, such as 
to a superficial observer exhibit the most striking 
resemblance to intelligence, upon closer inspection turn 
out to be evident proofs of the want of individual 
intelligence in animals. The brighter the light, the 
darker the shadows. 

The sanguine slavemakers afford us far better 
ground than the Amazons for assuming, that in their 
military expeditions individual intelligence comes in 
for a considerable share. Some scouts of F. sanguinea 
happening upon a nest of some slave species return 
with the news. As soon as the favorable moment 
for an expedition has arrived, they go ahead showing 
the way. Upon arriving at the hostile nest they gener- 
‘ally do not rush blindly to the attack, but institute a 
formal blockade; then, whilst one detachment impet- 
uously forces its way to the interior, others keep a 
sarcit watch on the outside and relieve the flying 


50 Chapter II. 


inhabitants of their larvae and pupae, which are the 
only objects the robbers have in view. On the part 
of the sanguineas this shows great cunning and looks 
very much like intelligence. If a troop of. apes at 
war with others were to surround the forest home of 
their foe and if a select squadron of the assailants were 
to penetrate into the woods, whilst the other part lying 
in ambush tried to capture the fugitives, how our 
modern evolutionists would be delighted with these 
apes! Such an argument for animal intelligence they 
would deem absolutely irrefutable, and they would 
no doubt allow this to be an “intelligent stratagem.” 
But sad to say, not apes but merely ants are skilled in 
such stratagems; yes, ants whose brain “can by no 
means compare with the brain of the higher animals!” 
If the development of the brain is the real cause of 
intelligence, then, of course, apes ought to be at least 
as intelligent as ants, or rather far more intelligent. 
In reality the reverse is the case, and thus, things 
look rather queer for modern evolutionism. 

Let us return to the military tactics of the sanguine 
slavemaking ants. One characteristic feature, that of 
reconnoitering the nest they wish to plunder, they have 
in common with the Amazons. With these latter ants, 
according to Forel’s observations, and my own, single 
individuals are wont to set out to investigate the site 
of a slave nest, and thus frequently enable the whole 
army of Amazons to advance in serried columns over 
a distance of thirty yards or more almost in a straight 
line to the place they had marked out. This surprising 
fact repeatedly observed by Forel and by myself can 
not be explained in any other than the above-men- 


Wars and Slavery in the Animal Kingdom. 51 


tioned way. With the Amazons, however, this system 
of reconnoitering cannot possibly have anything to do 
with individual intelligence, but only with instinctive 
sense faculties. This we have sufficiently proved - 
before. Hence, the necessary conclusion is, that also 
in the case of the sanguine slavemakers, instinct aided 
by sensitive experience will suffice to explain the same 
fact. Therefore, to postulate intelligence proper for 
such an explanation would be arbitrary humaniza- 
tion of the brute. 

The other seemingly very intelligent feature in the 
military tactics of F. sanguinea is their habit of setting 
out in smaller detachments more or less independent, 
and uniting only when one of the bands has some- 
where met with stronger resistance. Since most 
colonies of the negro ant (F. fusca), which is the 
ordinary object of their slave hunting expeditions, are 
not very populous, and as the inmates generally take 
to their heels at the first attack of the sanguineas, the 
latter’s tactics of dividing their forces is evidently 
appropriate to the usual conditions. Yet, if the assault 
is directed against an unusually populous and well 
defended nest of F. fusca, or against a large nest of 
the far more warlike F. rufibarbis, the same tactics 
frequently prove very disastrous to a considerable part 
of the assailing sanguineas. The first troop of the 
marauders venturing too close to the hostile nest is 
attacked and overpowered by the defenders, and sus- 
tains great losses, before any of the robbers are able 
to hurry back to call for assistance. If in the military 
tactics of F. sanguinea there were any question of intel- 
ligence or rational deliberation on the part of the single 


52 Chapter II. 


individuals, they would surely show the prudence and 
precaution of previously exploring more accurately 
the forces of the foe they want to attack. Thus, they 
would not dare an assault upon stronger slave nests, 
until a greater number of forces were collected; then 
they would, like the Amazons, fall upon the hostile 
nest in compact masses of many hundreds or thousands 
at a time, and would take the hostile position by storm 
without any considerable loss. Why does such a 
change never occur in the tactics of the sanguine slave- 
makers? A colony of these robbers, which for many 
successive years has pillaged the slave nests of the 
neighborhood and has experienced the different resist- 
ance offered by different hostile colonies, could easily 
remember their respective strength and could regulate 
the manner of future attacks according to this knowl- 
edge. It would be all the easier for them to make an 
intelligent use of their former achievements and 
reverses, because the worker ants generally live for 
the space of at least two or even three years. And yet 
not a trace of all this can actually be found. F. 
sanguinea will forever cling to her wonted tactics of 
setting out in small, scattered bands, even if bloody 
failure should ever so often be the result. To an 
unprejudiced psychologist such facts bear sufficient 
evidence of the fact that the warfare of F. sanguinea 
as well as of Polyergus is guided merely by hereditary 
instincts, not by individual intelligence. Those tactics 
were not invented by the intelligence of the ants; 
otherwise the same intelligence of the ants would be 
able to perfect and to develop them. Yea more: the 
assumption of ant intelligence is contradictory to the 


Wars and Slavery in the Animal Kingdom. 53 


fact that those tactics are specifically constant, and are 
specifically the same throughout the entire territory 
inhabited by F. sanguinea. 

Dr. Smalian has tried to invalidate this conclusion 
also. Here is his objection.’ “It was totally wrong of 
Wasmann to demand,? that the sanguineas should 
change their tactics, instead of continually attacking 
in small troops and thus being easily overpowered by 
large troops of fusca or rufibarbis. This mode of war- 
fare is inborn, and therefore instinctive, no less than 
the pillaging habit itself.” 

Dr. Smalian is wrong in ‘believing that we had 
in reality demanded of F. sanguinea to change her 
hereditary, instinctive stratagems. Our demand was 
merely the well known method of argumentation ex 
absurdo, which the critic seems to have misunderstood. 
In the supposition assumed by Smalian, but rejected 
by us, that ants besides their instinct possess also a 
certain degree of genuine intelligence, it is perfectly 
justifiable to demand that this intelligence should also 
be manifested and displayed. If their tactics are inborn 
only as to their outlines, this manifestation ought 
necessarily to consist in changing them intelligently 
according to circumstances, and consequently in their 
gradual perfection. But there is no trace of any such 
advancement towards perfection, and therefore we are 
right in concluding: These red marauding ants have 
only instinct, not intelligence. This mode of argu- 
‘mentation cannot seriously be styled “totally wrong.” 

Wherever the sanguine slavemakers live, they will 


1)°L.¢.,. p. 41. 
2) “Die zusammengesetzten Nester,” etc., p. 203. 


54 Chapter Il. 


follow the habit of invading nests of certain smaller 
species of Formica, and of rearing the robbed worker 
pupae, partly at least, as auxiliary ants for their own 
colony. It is, moreover, a constant characteristic of 
F, sanguinea to have rather a small number of slaves, 
if compared with those of the Amazons. . With these 
latter the slaves are far more numerous than the 
masters, with the former it is the reverse. Likewise, 
the specific military tactics are everywhere equally 
constant with both ant species. From the Alps to 
England and Scandinavia, from Holland to the 
Caucasus, F. sanguinea nowhere changes her habits 
and customs. Even her North American sub-species 
(rubicunda Em.) shows the same instinct of slave- 
making, and this in the same specific form. The only 
difference is, that one of the two European slave 
species, F. fusca, is represented in the North American 
rubicunda colonies by a closely allied variety, namely 
by F. subsericea.*_ Since the separation of North 
America from Europe was completed in the Tertiary 
age, the enslaving habits of the sanguineas and their 
military tactics must have been essentially the same in 
the Tertiary as they are today. This is the most natural 
explanation for the specific uniformity of that instinct 
in the different parts of the globe. One thing, how- 
ever, is certain: if the impulse of slavemaking and the 
specific military tactics of F. sanguinea were due to 
the intelligence of the ants, or if they were even in the 
slightest degree dependent on it, such a specific uni- 
formity existing for thousands of years would be 
utterly inconceivable. 


1) See Wasmann, ‘“‘Kritisches Verzeichniss der myrmekophilen und 
‘termitophilen Arthropoden” (1894), p. 163 ff. 


Wars and Slavery in the Animal Kingdom's 55 


3. The Pretended “Automatism” in the Psychic 
Life of Ants. 


Animal intelligence, therefore, has no part either 
in the slavemaking expeditions of ants, or in their 
military tactics. Yet, the application of these instincts 
is not mathematically uniform, They are influenced 
and governed by the changeable sensitive perceptions 
and individual conditions of the single ants, and thus 
great variability exists within specified limits. Those 
animal psychologists who, in contradistinction to the 
higher animals, call ants mere “instinct automatons,” 
or even mere “reflex machines,” are asked to consider 
that the instincts of ants are neither more nor less 
“automatic” than those of dogs, apes and other verte- 
brates. Instances of ‘intelligence in the true sense of 
the term can be discovered with the latter as little as, 
and even much less than, with ants. Various differ- 
ences, however, of individual character, and of. indi- 
vidual action, determined by different sense perceptions 
and sense experiences, occur with ants as well as with 
the higher mammals. 

On turning over the stone or the piece of sod 
covering a middlesized nest of F. sanguinea, and thus 
suddenly exposing the interior to the light, we perceive 
all the inhabitants in tumultuous excitement. Part of 
the ants furiously biting and ejecting poison attack 
the invader; others take care of their imperilled off- 
‘spring and in haste carry down the eggs, larvae and 
pupae to the lower chambers of the nest; other indi- 
viduals of the same colony seem destitute of the 
chivalrous spirit of their race for the defense of 


56 Chapter II. 


country and escape under sheltering grass tufts or 
clods of earth; sometimes, even, in the midst of her 
fighting, rescuing or fleeing comrades, a sanguinea 
presses herself to the soil motionless and, though 
mostly for a short time, has recourse to the instinctive 
trick of “feigning death ;” in opposition to these, other 
sanguineas, finally, seem to be seized by a strange 
mixture of courage and fear, by a sort of impotent 
rage: not venturing to attack the real foe, they vent 
their spite against other objects; with sprawling feet 
they crawl along the ground, and with their heads bent 
down they furiously bite the sand or stalks of heather, 
attacking everything, in fact, but the finger of the 
great human monster that robs their nest of Lome- 
chusast and other favorite guests. Such scenes as the 
one just described I have observed hundreds of times, 
and am so accustomed to them, that I find them quite 
natural; nevertheless, they are of the utmost import- 
ance for comparing the psychic faculties of ants and 
those of the higher animals. Packs of wolves or hordes 
of apes on similar occasions could display no greater 
variability of individual character and of individual 
action, than such a colony of sanguineas. Yet, ants, 
we are told, are “instinct automatons,’ and apes or 
wolves are not! 


1) In the colonies of the North American subspecies of sanguinea, 
‘F. rubicunda, the European Lomechusa strumosa is represented by an 
allied species, Xenodusa cava. Rev. H. Muckermann, S. J., of Prairie 
du Chien (Wisconsin) has recently found also the curious pseudogyne 
ant form, which is due to the education of the larvae of Lomechusini 
by the ants, in the colonies of F. rubicunda. See Wasmann, ‘Neue 
Bestaetigungen der Lomechusa-Pseudogynentheorie” (Verhandl. der 
Beutsch. Zool. Gesellsch. 1902, p. 98-108 and PI. II.). We shall give 
the figure of Xenodusa later on, opposite p. 181. 


Wars and Slavery in the Animal Kingdom. 57 


A dog biting the stone thrown at him, in his blind 
rage acts just as ‘‘automatically” as a sanguinea which 
vents her fury on the edges of a glass tube, so that you 
can hear the grating noise made by her jaws. And if 
certain individuals of an ant colony acquire through 
their sense-experience special dispositsons and char- 
acteristics, which distinguish them from other individ- 
uals of the same colony, then they act “automatically” 
as little as dogs or apes, or other higher mammals do. 
Some remarkable instances of this may find a place 
here. 

In the observation nest of F. sanguinea described 
on page 23, some beetles called Dinarda dentata, which 
I introduced, had at first been received as usual without 
difficulty as indifferently tolerated guests, and had 
even propagated in the nest. But several times I put in 
a little larger Dinarda species (D. Maerkelii), whose 
usual host is F. rufa, and when finally some small 
sanguineas and their slaves had succeeded in seizing 
and killing this beetle, which, as a rule, is unassailable 
owing to its wedge-shaped body offering scarcely any 
point of attack,t then a number of ants of this colony 
gradually took a liking to catching Dinardas, which 
liking proved disastrous also to the smaller Dinarda 
dentata. Not all the individuals of the different ant 
species of that colony have acquired this strange 
passion. Among twelve workers of F. sanguinea 
which I put from this observation nest into a smaller 
_ experimenting nest together with seven Dinarda 

1) See Wasmann, “Dinarda-Arten oder -Rassen,” in Wien. Entom. 
Ztg., 1896, 4th and 5th issue, and ‘‘Die Myrmekophilen und Termi- 


tophilen,” p. 435 (Extr. du Compte rendu du troisiéme Congrés intern, 
de Zool. Leyden, 1896). 


58 Chapter II, 


Maerkelii, there was but one Dinarda hunter. Whilst 
the rest remained perfectly passive towards the 
Dinarda, this one ant immediately began an active 
hunt. Had I not soon removed her from the small 
nest, she would probably have aroused in her com- 
panions the inStinct of imitation for a similar persecu- 
tion, a fact which I have often observed. But by 
remoying this passionate hunter, I preserved friendly 
relations between the other ants in the same experi- 
menting nest (11 sanguineas, 2 rufibarbis, 2 fusca) 
and the Dinarda Maerkelu. In the greater observation 
nest, from which I had taken these individuals, the 
Dinarda hunt, which had begun with the killing of 
Dinarda Maerkelu in March 1896, continued against 
D. dentata until November of the same year, when the 
ants gradually returned, but only for a short time, to 
their former toleration of these guests. The resuming 
of experiments in the following spring resulted finally 
in the complete extermination of all the Dinardas in 
that observation nest. During the following six years, 
I never succeeded in securing the existence of even a 
single D. dentata in that nest, although in nature this 
beetle is indifferently tolerated in all sanguinea nests! 
The psychological importance of these phenomena has 
been pointed out already in our discussion on the 
different forms of learning in the animal kingdom. 
Another strange fact quite irreconcilable with the 
“blind automatism” of instinct is the behavior of ants 
regarding the number of Dinardas and other beetles, 





1) “Instinct and Intelligence,” etc. (Herder, St. Louis, Mo., 1903), 
p. 157. Also “Die psychischen Faehigkeiten der Ameisen” (Stuttgart, 
1899), pp. 84, 88, 93, 


Wars and Slavery in the Animal Kingdom. 59 


which I introduced into that observation nest during 
my experiments on the international relations’ of ant 
guests. If previously no Dinarda had been present for 
some time, one or two of D. dentata were often quietly 
received and tolerated for weeks. But as soon as I 
would add some more beetles of this species, the perse- 
cution began, at first against the newcomers, and 
finally, the passion of hunting having been aroused, 
also against the old ones, until all were seized and 
devoured. Thus the ants seem to have been led to this 
persecution not so much by the perception of one beetle 
by itself, but rather by that of their increasing number. 
It was this that aroused their hostility. 

The sanguineas of the same observation nest once 
manifested unmistakably that they desired only a 
limited number of their genuine guest Lomechusa 
strumosa, otherwise so dear to them. In September, 
1898, I had captured in a sanguinea colony? 116 beetles 
of that species and placed 30 of them in my observa- 
tion nest. After a few weeks, however, 19 of them 
had been driven out of the main nest and were con- 
fined in the empty annex, where they all perished, 
being isolated from their nurses, whilst the remaining 
11 were for the whole winter carefully tended by their 
hosts and very often fed and licked by them. The 
ants, therefore, had not changed their behavior toward 


1) “International relations’ I term the relation of ant guests to 
different colonies and different species of ants, which are hostile to 


. @ach other. 


2) No. 191 of my statistical map of the sanguinea colonies near 
Exaten. In treating about the education of the Lomechusa larvae by 
F. sanguinea, I shall communicate more detailed observations regarding 
this colony (in the last chapter, number 3, “Adoption Instincts in the 
Animal Kingdom’’). 


60 Chapter II. 


the Lomechusas as such, but merely toward their 
excessive number, which became disagreeable to them. 
Perhaps they were unable to feed so many beetles and 
had therefore expelled half of them. At any rate, 
such phenomena prove to a certainty that ants are 
not to be regarded as mere “instinct automatons” or 
“reflex machines.” We must ascribe to them sensile 
mental faculties, which by way of different percep- 
tions and representations cause great variability in the 
display of their instinctive impulses. But, beyond this, 
nothing is required to explain satisfactorily the psychic 
life of the vertebrates. Hence, there is no need of 
“animal intelligence,” neither in the case of ants nor 
in that of the higher animals. 

A beautiful instance of how sensile experiences of 
ants lead them to acquire certain individual peculiar- 
ities of character, I witnessed in the case of a F. rufi- 
barbis of the same mixed colony. She was a worker, 
easily distinguishable from the others by her small size. 
She used to visit regularly the glass bulb of the feed- 
ing tube (see diagram, p. 23), where she would lick 
the honey or sugar in order to supply the other ants 
in the main nest from the sweet juice stored up in 
her crop. Although F. rufibarbis belongs to a very 
irritable and pugnacious species, yet this ant had 
gradually become so tame that she would allow her- 
self to be fed from my hand. As soon as I removed 
the cork of the glass bulb, she would come out and 
look for food on the outside. I would then present 
to her a needle dipped in honey. At first she darted 
back, but after a few seconds of hesitation she would 
approach, examine the needle with her feelers and 


Wars and Slavery in the Animal Kingdom. 61 


lick off the honey. Later on I placed the honey on my 
finger. The ant had already become so tame that 
she was not in the least disturbed by the odor of my 
finger, whereas others would have been provoked to 
a fight or would have been greatly alarmed. She 
would quietly lick off the honey and then, without 
resisting or trying to flee, allow herself to be seized 
with a pincette by one of her legs and placed back in 
her nest. This goes to prove that ants also are tama- 
ble in spite of their excitable nature. The tamable- 
ness of ants, like that of higher animals, is due to their 
possessing the powers of sensitive perception and imag- 
ination, upon which the intelligence of man acts to 
accomplish his purpose. 

Against this parallelism an objection was raised 
by Mr. Bethe.t To tame an ant, he says, takes 
weeks and months; but a dog may be tamed in a few 
days; therefore the above mentioned fact presents no 
proof of the existence of psychic faculties in ants! 
Whether Mr. Bethe will succeed each time in taming 
a vicious dog within a few days, is rather doubtful. 
Nor is it at all true, that it takes several weeks or 
months to tame an ant, e. g., a F. fusca or rufibarbis, 
which are especially suitable for such experiments. 
It is but required to mark a certain individual which 
comes regularly to the feeding tube. If you are very 
careful not to frighten the animal, it is possible to 
train it in a few days, in the manner described above. 
' But if you wish to reckon the time needed for taming 
an ant by beginning with the day on which she was 


1) “Duerfen wir den Ameisen und Bienen psychische Qualitaeten 
zuschreiben?”? (Bonn, 1898), p. 23. 


62 Chapter II. 


first deprived of her freedom and placed in the arti- 
ficial nest, then the same method of calculation must 
be applied in the case of the dog. Mr. Bethe should 
not, therefore, take for his experiments a domesticated 
dog, but he would have to operate upon a newly- 
captured animal of the wild dog species. Then let 
us see, which would take longer, to tame a wild dog 
or to tame a wild ant! 

There are several other interesting analogies 
between the psychic life of ants and that of dogs. 
A small dog, as long as he is in the company of his 
master or of some stronger comrade, will not be afraid 
to meet a rival, whom otherwise he would try to avoid. 
The same is the case with the small black negro ants 
(F. fusca) when they are in company with sanguineas. 
In their own colonies they are generally cowards. 
As soon as their nest is disturbed, they flee and try to 
hide their young, but when they are slaves in colonies 
of F. sanguinea, they are the bravest defenders of the 
mixed colony, as I have often experienced to my cost. 
Just as in the mixed colony of F. sanguinea the 
instinctive courage of F. fusca, which is otherwise so 
cowardly, is to be explained psychologically from their 
perception of the great number of valiant companions 
and their consequent sense of solidarity, without sup- 
posing any reasonable deliberation on their part, so 
also are the different degrees of courage found in 
different colonies of the sanguine slavemakers to be 
accounted for. If a numerous population inhabits a 
rotten fir stump, on the surface of which we find some 
of the ants running about, a gentle kick will at once 
call forth a whole army ready for the fray. In a 


Wars and Slavery in the Animal Kingdom. 63 


moment the whole surface of the stump is covered 
with thousands of ants furiously hurrying to and fro. 
But, if the colony is weak, the same kick, which at 
other times calls forth an army, will have the con- 
trary effect. The ants which just before were running 
about the surface, disappear through the entrances of 
the nest as if by magic, and deathlike quiet succeeds, 
If in ants this appropriate estimation of the strength 
of their own colony is characterized as instinctive,— 
and this is, no doubt, the only correct expression,— 
then, similar occurrences among higher animals should 
also be credited to instinct and not to intelligence.’ 
Yet, the courage of individual ants in a colony is 
dependent not only on the perception of the great 
number and courage of their comrades. In populous 
colonies also of warlike species such as F. sanguinea, 
there will always appear considerable differences in 
individual courage, as we have shown above. Some- 
times even single, isolated individuals make head 
against a numerous foe. Such an example of “hero- 
ism” scarcely ever equaled by dogs, lions, and tigers, 
was once witnessed by Rothney? in Bengal. A mid- 
dle-sized worker of a large black ant species (Cam- 


1) Forel (Un apergu de Psychologie comparée [1896], p. 25) men- 
tions an instance, where a very strong colony of Camponotus ligni- 
perdus, when fighting with F. pratensis, showed a more warlike spirit 
-than is the custom with the ordinary colonies of that species. We fully 
agree with Forel if he infers therefrom the existence of “plasticity” 
in the psychic faculties in ants. Our own observations mentioned 
‘above prove the very same. Yet, in considering this plasticity of the 
sensitive powers of cognition and appetite to be essentially identical 
with human intelligence, he i$ entirely wrong, as we have shown in a 
former publication, ‘Instinct and Intelligence in the Animal Kingdom.” 

2) “Notes on Indian Ants,” p. 349 (Transact. Entom. Soc. Lon- 
don, 1889). 


64 Chapter IT, 


ponotus compressus) attacked quite alone a whole 
colony of small red ants (Solenopsis geminata). 
Without moving from her place she remained before 
the entrance of the Solenopsis nest from 4:30 p. m. 
until night, seized the ants with her jaws as they came 
out, and bit them in two. At last she was overpowered 
by superior numbers, and after having killed 150 or 
200 of her foes she paid for her temerity with her 
life. We leave it to the modern worshippers of animal 
intelligence to raise a monument to the memory of 
this insect Leonidas. 

We need not, however, go as far as Bengal to find 
‘such examples of “heroism” of single ants. There 
are plenty of them in the heaths of Germany and 
Holland, and also in North America. During hot 
weather strolling workers of sanguinea will frequently 
enter into a fierce battle with colonies of Lasius niger 
or Tetramorium caespitum living in the neighborhood, 
till at length, when too many of their opponents have 
clung to their legs, they fall victims of their own fool- 
hardiness. A scene, more harmless indeed, yet psy- 
chologically not less remarkable, I observed near 
Exaten on the afternoon of August 15, 1894. A large 
worker of F. sanguinea amused herself for a quarter 
of an hour by blockading all alone, a colony of the 
small, red stinging ants (Myrmica scabrinodis). She 
lurked about the entrance, seized by the neck one red 
ant after another, as they came out, carried them quickly 
to a distance of several inches, and dropped them in 
order to be back again at once at the entrance to seize 
the next customer. The Myrmicas scarcely attempted . 
any resistance, although several dozens of them were 


Wars and Slavery in the Animal Kingdom. 65 


_in and around the entrance. Only one or two tried to 
get hold of the intruder by one of her legs, but with- 
out success. Of course, their horny (chitine) armor 
protected them sufficiently against the jaws of the 
sanguinea; all the more, as the latter did not allow 
herself time to pay special attention to her single foes. 
It was exceedingly amusing to see the indefatigable 
zeal and hurry of the large ant getting hold of the 
small ants one after another and carrying them away, 
after which they slowly crawled home again. It is 
hard indeed to say what induced the sanguinea to 
engage in this odd kind of skirmish. Perhaps it was 
mere love of fighting. That she released the single 
Myrmicas so soon, might be explained by her fear 
of the sting, with which these ants are provided. Yet, 
it is more probable, that the marauder took a fancy 
to the entrance of the Myrmica nest, and on this 
account tried to expropriate the inhabitants. Of 
course, her labor was no more successful than that of 
Sisyphus, because she did not carry the ants further 
than a few inches from the nest, and, besides, the 
number of ants coming out of the nest had no end; 
but this did not seem to affect her in the least. 

It would be ridiculous, arbitrarily to humanize 
such instances and to suppose all possible kinds of 
“intelligent purposes” on the part of the ant, as is 
customary with popular psychology. Yet, on the other 
hand, it cannot be denied, that “mechanical autom- 
atism” of instinct will never explain them. The only 
satisfactory solution psychology can give, is to 
ascribe to ants sensitive powers of cognition and appe- 


tite, which, under the influence of exterior sense per- 
5 


66 Chapter II. 


ceptions and individual dispositions, are the principle, 
from which these various, spontaneous activities result, 
with no difference as to whether the actions are per- 
formed by ants, or by dogs and apes. 

The phenomena described above may be classed 
among the “sports” or “games” of animals, as Groos? 
terms them. The facts recorded deserve these names 
perhaps just as well as the sports and games of the 
higher animals; only, it is generally much more diffi- 
cult to ascertain the nature of given facts in the case 
of ants. Among the heaps of ants that gather on the 
surface of the ant-hills of F. rufa and pratensis, as also 
in my artificial nest containing sanguineas when 
exposed in spring to the warm rays of the sun, I have 
repeatedly observed instances of harmless wrestling, 
beginning with and accompanied by lively and playful 
movements of the feelers. This behavior of the ants 
seems to be due to a resuscitation of their powers and 
also to an excess of muscular energy after the winter’s 
rest. ; 

Forel (Fourmis de la Suisse p. 367) has made 
similar observations with F. pratensis, and Huber? with 
F, rufa and pratensis. I cannot consider these games 


1) “Die Spiele der Tiere” (1896), pp. 125 and 135. By the way, 
Groos here and elsewhere was too confident in trusting the authority 
of Buechner, who has not unfrequently misrepresented Huber’s and 
Forel’s observations to suit his own purposes of humanizing the brute. 
Forel, in the Etudes myrmécologiques, has expressly protested against 
Buechner’s misrepresentations of his observations. The book of Groos 
contains in general a great many statements of doubtful value, in spite 
of the critical standpoint from which the author maintains to view the 
facts. 

2) Since Huber in his “Recherches,” p. 151, does not say whether 
he means the fourmi fauve 4 dos rouge or that @ dos noir, we are 
hardly able to decide which ant it is. 


Wars and Slavery in the Animal Kingdom. 67 


of ants as an “evident reaction against instinct,” as 
Forel did, at least formerly, in his book mentioned 
above. They are instinctive no less than the romps 
and scuffles of young cubs, or the frolics of lambkins. 
They are due, I suppose, to the natural impulse of 
exercising the muscles, which is pleasurable to animals 
as well as to human beings. At any rate it would be 
wrong to ascribe them to intelligence, either in ants, 
or in the higher animals. 

The erroneous doctrines about the “absolute blind- 
ness” and the “mechanical automatism” of instinct, 
which are still current, have done a great deal towards 
making animal intelligence appear almost indispen- 
sable. Assuming instinct to be a mere reflex mechan- 
ism, it was of course impossible not to declare as “‘intel- 
ligent activities” all the manifestations of the instinct- 
ive faculties, which are determined and influenced by 
individual sense perceptions and sense experiences. 
This, however, as we have shown above,' is an alto- 
gether uncritical procedure. The study of ant life 
enables us to determine more correctly the nature of 
animal instinct, this being the case especially with 
F. sanguinea, which in point of so-called “intelligence” 
may fairly rank with the highest vertebrates. 


4, Slavery among the Sanguine Slavemakers. 


The custom prevalent among these red ants of rob- 
bing and rearing slave pupae, is of course entirely 
instinctive. Even our scientific opponents acknowledge 
that it would be nonsense to explain it as an intelligent 


1) “Instinct and Intelligence,” etc. (Herder, St. Louis, Mo.), Chap. 
II and III. : 


68 Chapter II. 


invention of some colony of sanguineas, transmitted 
by inheritance to all the descendants of the species. 
Forel, Emery and Smalian fully agree with our tren- 
chant condemnation' of Buechner’s manner of human-: 
izing the “slavery” of ants. It does not seem impos- 
sible, however, that for the actuation of this instinct 
there should be needed special psychic impulses pro- 
duced in the young ants by the example and the feeler 
language of their older companions. Yet, this assump- 
tion is scarcely probable; for the formation of new 
colonies is undertaken, as a general rule, by single 
impregnated females; but the females of F. sanguinea 
are devoid of the enslaving instinct, and cannot, there- 
fore, induce others to manifest it. Yet, since it is the 
general opinion, that tradition and instruction aid the 
exercise of the social instincts in these insects, and that 
the high perfection of their community life receives 
thereby its full explanation, we will examine whether 
in view of the facts this opinion is still tenable. 

It is true, in ant communities the instinct of imita- 
tion plays a great part, as we may gather from several 
observations recorded above. By the example and the 
taps of the feelers of their older comrades the younger 
ants are often induced to actions, which otherwise, 
at least under the same circumstances, they would not 
have performed. In this regard, as in fact in the other 
salient features of the psychic life of animals, ants and 
the higher animals agree in all the essentials; for, 
in the latter also the so-called lessons given to the 
young by their parents consist only in exciting instinct- 
ively in the young the faculty of imitation by the exam- 


1) “Die zusammengesetzten Nester,” p. 182. 


Wars and Slavery in the Animal Kingdom. 69 


ple of their parents. If we interpret “tradition and 
instruction” in this sense, it must be acknowledged 
that they aid in the exercise of the hereditary instincts 
both in ants and in the higher animals. But, on the 
other hand, it is equally obvious, that in this case the 
terms “tradition” and “instruction” mean something 
very different from what modern animal psychologists 
wish to insinuate; for, in our case, they do not imply 
any intelligent communication of knowledge, but only 
the instinctive excitation of the imitative faculty. 

But in the communities of social insects not even 
the encouraging example of the older companions is 
necessary for the first actuation of the young workers’ 
instincts. We have ascertained by experiments, that 
precisely the most remarkable and apparently most 
intelligent habits of the sanguine slavemakers, namely 
their rearing of slaves and the hospitable care bestowed 
by them on the beetle Lomechusa strumosa, are merely 
hereditary instincts, for the exercise of which no kind 
of “instruction” on the part of the older ants is 
needed.’ To prove this we formed a special colony of 
“self-taught” young workers of F. sanguinea, by plac- 
ing in a glass filled with a sufficient quantity of earth 
a number of ants that were newly developed from their 
cocoons in my artificial nest. These self-taught ants 
not only performed all the works required for building 
their nest, just as the other individuals of their species, 
but. they also followed the very same line of conduct 
. in nursing their young and even in dealing with 
strange worker pupae which I introduced into their 


1) L. c., p. 202, and “Die internationalen Beziehungen von Lome- 
chusa strumosa,” in the “Biologisches Centralblatt,” XII (1892), 592, 


70 Chapter II. 


nest. The pupae of Lasius niger they would either 
devour or throw away, whereas those of F. rufibarbis 
were reared by them as auxiliary ants for their colony. 
A Lomechusa strumosa which IJ put in, was immedi- 
ately received like an old acquaintance, licked and fed, 
just as is the custom in the other colonies of F. san- 
guinea. In the face of such experiments the beautiful 
theory of tradition and instruction among ants van- 
ishes into thin air. 

That the older ants “lead their newly born com- 
rades about the nest and train them to a knowledge 
of domestic duties, especially in the care of larvae,” 
is a fable originated by Buechnert and unfortunately 
taken up on his authority even by Rorhnanes? and 
other modern animal psychologists. The truth is, that 
the newly developed ants are as yet the objects of spe- 
cial care and protection on the part of the others, as 
remarked already by Huber. Being as yet rather help- 
Jess, they are still, as it were, considered as “wards.” 

The same applies to bees. In their case also the 
instruction said to be given by the old workers is a 
mere fancy, arising in the brain of some anthropomor- 
phizing observer. Already Réaumur in his classical 
work Histoire des Insectes* remarks: “Scarcely have 
all the parts of the body of a young bee become suf- 
ficiently dry, scarcely is she able to move her wings, 
when she is already acquainted with everything she 
will have to do in the whole course of her life.” He 
goes on to relate a few observations showing, that 


1) “‘Geistesleben der Tiere,” p. 62. 
2) “Animal Intelligence”? (6th ed.), p. 59. : 
8) Tom. V. part II, mém. XI, p. 278. Amsterdam, 1741. 


Wars and Slavery in the Animal Kingdom. 71 


young bees from the very first day are as well able to 
use their instincts as are their seniors. Of late some 
experiments have been made by Kogevnikov' and 
Butkewitsch? on self-taught young bees. The results 
were practically the same as in the case of the self- 
taught ants. It was found that in the workers the 
building of combs and the nursing of the young, and 
in the queens the love of combat were hereditary 
instincts, utterly independent of experience and instruc- 
tion. Besides Charles Janet’s excellent observations 
on hornets* show, that social insects are ruled only by 
hereditary instincts, excited to their natural manifes- 
tation by the very first experiences of the young indi- 
vidual. The impulse of imitation with its various 
incitements is only a secondary factor. This is the 
truth regarding the captious shibboleth of “instruction 
and tradition” in insect communities. 

No doubt, therefore, is left as to the fact that the 
slavemaking habits and the military tactics of the san- 
guineas, just as the social life of ants in general, are 
due to instinct only, not to individual intelligence. 
Yet, this instinct is not an absolutely blind impulse, 
but is suitably modified according to the wants and 
purposes of a given colony. A blind impulse to rob 
and to rear slave pupae would be expected to impel 
sanguinea colonies to rob the more slaves, the stronger 
and more numerous they are themselves. In the most 
populous nests we ought to find the greatest number 


1) “Zur Frage vom Instinct,” in ‘Biolog. Centralbl.,” Vol. XVI 
(1896), No. 18, pp. 657-660. 

2) “Russisches Bienenzuchtblatt,” April,1896. | See Kogevnikov 1. c. 

8) “Mémoires de la Société Zoologique de France.” T. VIII 
(1895). . 


72 Chapter II. 


of auxiliary ants. In reality the very reverse happens. 
The most populous sanguinea colonies do not contain 
the relatively greatest but the relatively smallest num- 
ber of slaves. We formerly (in “Die Zusammengesetz- 
ten Nester” p. 50) alluded to this fact, which shall now 
be explained and proved at greater length. In order 
to show the connection between the rearing of Lome- 
chusa strumosa in. the sanguinea nests and the educa- 
tion of a strange, crippled kind of workers, the 
so-called pseudo-females or pseudogynes,? I drew up 
an accurate statistical map of the sanguinea colonies 
in the neighborhood of Exaten. It comprises 410 
colonies with more than 2,000 nests.? Regarding the 
number of slaves, the statistics showed that in most 
colonies the masters were from three to six times more 
numerous than the slaves. The most populous colonies 
contain scarcely 50 to 100 slaves, sometimes even less 
or none at all. In the middle-sized or weaker colonies, 
however, the absolute number of slaves amounts in 
most cases to several hundred. The average propor- 
tion of masters and slaves in the most populous 
colonies is from 100:1 to 10:1, in the middle-sized 
and weak colonies, however, from 3:1 to 1:1. Nor 
are these the ultimate limits assigned to the number 
of slaves found in the nests of these ants. In May, 
1890, and from 1896 to 1898 I found near Exaten 
several strong sanguinea colonies without any slaves.* 


1) “Die ergatogynen Formen bei den Ameisen und ihre Erklae- 
rung,” in “Biolog. Centralbl.,” Vol. XV (1895), Nos. 16 and 17. 

2) A colony of F. sanguinea not unfrequently embraces several 
nests, often one or more metres distant from one another, inhabited 
all at the same time or alternately. 

3) To similar colonies of F. sanguinea of the race rubicunda in 
North America we must probably refer the F. sanguinea race aserva 
of Forel, who described it lately from Toronto (Canada), (Ann. Soc. 
Ent. Belg. XLV, 1901, p. 395). 


Wars and Slavery in the Animal Kingdom. 73 


A similar colony I recently detected near Luxemburg, 
1904. On May 23, 1889, I met with the opposite 
extreme, namely, a very weak sanguinea colony, in 
which the slaves were about twenty times more 
numerous than their masters. These extreme cases 
are, however, very rare. Besides, it is plain that the 
number of slaves in different colonies changes every 
year; and lastly, the number of slaves in the nests of 
the sanguine slavemakers depends also on special, 
local circumstances. Where slave nests are very 
numerous, e. g., in groves of birches and oaks, there 
more slaves will be found in the sanguinea nests than 
on the open heath, where fusca nests are very scarce. 
Yet, ceteris paribus, we find the constant law, that in a 
given sanguinea nest, the number of slaves and that of 
masters is not in direct but in inverse proportion. My 
observations of sanguinea nests in Dutch Limburg, 
Rhineland, Vorarlberg, Bohemia and Luxemburg, 
everywhere confirmed this law. 

How is this remarkable difference between the col- 
onies of sanguineas and of Amazons to be explained? 
The latter possess the more slaves, the more populous 
the colony; with the former we meet the reverse. This 
difference is explained by the fact that F. sanguinea 
is not, like the Amazons, essentially dependent on her 
slaves, but rather regards them, as it were, as a 
secondary complement of her own communities. The 
sanguineas rob and rear only as many slave pupae, as 
' is suitable for their colonies. Weaker colonies thus 
feel greater need of supplementing their own deficiency 
by adding auxiliary forces, whereas stronger colonies 
do not feel the same necessity; so they regulate their 


74 Chapter II. 


action according to their perception of this deficiency. 
It is true, the smaller number of slaves in more popu- 
lous sanguinea nests may partly be due to another 
circumstance, viz., that in the latter a greater per- 
centage of robbed pupae is eaten than in smaller nests. 
Nevertheless, this circumstance is far from explaining, 
why in the weakest colonies of this marauding ant the 
number of slaves even exceeds that of the masters. 
The only way of accounting for this fact is to assume 
that these colonies try to strengthen their forces by 
the greatest possible number of auxiliaries. 

To perceive this necessity of increasing their 
numbers does not go beyond the limits of the 
instinctive powers of ants. A very interesting case of 
this kind was observed by me both in the summer and 
the fall of 1898 in my artificial nest of F. sanguinea, 
already repeatedly referred to. During my absence, 
extending over several weeks of July and August, the 
nest had been badly cared for and had repeatedly dried 
up; consequently many workers of sanguinea and the 
greater number of the old slaves had perished. Mean- 
while new auxiliary ants (F. rufibarbis) had been 
reared from cocoons which I had given to the 
sanguineas. Now, I observed in the course of Sep- 
tember, that new clusters of eggs, laid by the two 
queens, made their appearance in the nest, and that a 
number of larvae were reared, some of which were 
conspicuous for their rapid development. This is an 
exceptional case, generally not occurring in sanguinea 
colonies during autumn. In October and the first part 
of November I was absent again. On my return I 
found, to my great surprise, that in place of the two 


Wars and Slavery in the Animal Kingdom. 75 


wingless queens there were now four of them in the 
nest. This shows that the ants had reared two new 
queens from the eggs which had been laid in the fall.* 
The mortality prevalent in their colony during summer 
must have caused this exceptional conduct. On duly 
considering such facts, it can hardly be found strange 
that sanguineas try to make up for the deficiency of 
their numbers by robbing and rearing the pupae of 
other ants; for the enslaving habit serves them as a 
secondary means of increasing their colonies. 

The manner, too, in which the sanguineas make 
their choice between the different castes of alien 
Formica pupae, confirms the above explanation; for 
this phenomenon is understood only by referring it to 
their desire to augment the forces of their colonies. 
It is evident that the rearing of slave pupae is not 
governed by a “blind instinct of education,” inducing | 
the ants to transfer all their care to the strange brood; 
for they consume the male and female pupae of 
strangers, or kill the sexual ants as soon as they leave 
their cocoons, whereas they adopt, at least partly, those 
pupae of strangers, which will develop into workers. 
They distinguish, therefore, by olfactory perceptions 
between their own pupae and those of strangers, and 
they likewise distinguish between the different castes 
of the latter. Hence, we are compelled to maintain: 
the sanguine slavemakers do not rear the worker 
pupae of strangers, because they are unable to distin- 
guish them from their own, but because they aim at 
augmenting the number of their own workers by 
~~ 1) The two old queens were still living in the spring of 1900, and 
laid eggs in spite of their age of almost ten years. One of them died 


last year (1903), thirteen years old. The surviving queen (fourteen 
years old) I set at liberty with the rest of the workers in spring, 1904. 


76 Chapter II. 


adopting strangers as auxiliaries. Of course, they are 
far from having an intelligent knowledge of this pur- 
pose. It suffices that, on account of the actual need of 
workers, the instinctive impulse of the ants to bring up 
fresh workers is aroused with greater intensity and, 
for this reason, extends to other Formica cocoons. 
And this is the only explanation admissible, for we 
have proved above, that with F. sanguinea slave- 
holding is not due to experience or instruction, but to 
hereditary instincts. 

Is there anything in the social life of higher 
animals, which can rival this strange phenomenon? 
We know of nothing. If there had been, then Darwin, 
Ziegler and other evolutionists would not have failed 
to turn it to account, and to appeal to it as convincing 
proof of the “quasi-human intelligence” of higher 
animals; for, if an association of animals perceives 
the necessity of increasing its strength by adopting 
auxiliary forces, and under the influence of this per- 
ception actually adopts them, then this action proceeds 
from a motive originating in sensitive experience, and 
is therefore intelligent, at least according to modern 
animal psychology. Nevertheless, ants are said to be 
“instinct automatons,’ but higher animals are not! 
And this again shows, how utterly untenable, on the 
one hand, is the modern notion of intelligence, and, 
on the other, how foolish the attempt to place the 
“intelligence” of the higher animals on a far higher 
level than that of ants. 

Bethe,’ indeed, has of late made an attempt to 
explain, in a very simple manner, the proportion 


1) “Duerfen wir den Ameisen und Bienen psychische Qualitaeten 
zuschreiben?” p. 69. 


Wars and Slavery in the Animal Kingdom. 77 


between the number of masters and of slaves found 
in colonies of F. sanguinea. “The correlation,” he 
says, “existing between the numbers of masters and 
slaves is as little owing to psychic processes, as the 
numerical correlation existing between mice and 
buzzards, or between certain butterflies and cuckoos.” 
Yet, it is hard to see what is proved by this compari- 
son, unless it be the very contrary of what Bethe 
pretends to prove; for the more mice there are, the 
more buzzards will come to the spot, and the more 
butterflies, the more cuckoos; however, in the case of 
sanguinea colonies just the reverse takes place, namely, 
the more masters there are, the fewer slaves they have 
in their colonies! 

As the sanguineas accommodate themselves to 
given circumstances regarding the number of their 
slaves, so also regarding their species. Their favorite 
slave species is F. fusca. This black ant is found as 
auxiliary in the greater number of the above men- 
tioned 410 sanguinea colonies near Exaten. In 25 
colonies the place of F. fusca is taken by a different 
species, viz., F. rufibarbis; 17 colonies have both 
species. Near Feldkirch, in Vorarlberg (Austria), I 
found side by side with colonies which had the above 
mentioned slaves, others with F. cinerea, or with F. 
fusca and cinerea. The latter species does not occur in 
Dutch Limburg, and for this reason no cinereas are 
found there as slaves in sanguinea nests. Yet, the fact 
that the sanguineas occasionally invade weak colonies 
of the large hill ants (F. rufa and pratensis) to rob 
their pupae and to rear them as auxiliaries, proves 
that their “blind instinct” does not force them to rob 


78 Chapter II. 


automatically a certain, fixed species of slaves. In 
May, 1890, I found near Exaten such a “natural, 
abnormally mixed” sanguinea colony containing, 
besides F. fusca, a considerable number of F. rufa. 
This colony has since disappeared; for on my return 
after a two years’ absence I looked for it in vain. 
Since 1895 I found in the same neighborhood of 
Exaten four other natural, abnormally mixed colonies 
of sanguineas. One of them (col. No. 66) had only 
F. pratensis as slaves; the second (col. No. 105) 
F. rufo-pratensis, a variety intermediate between rufa 
and pratensis, with F. fusca; the other two had (col. 
Nos. 84 and 247) F. pratensis and fusca. Three of 
these colonies, therefore, possess besides the ordinary 
species of slaves, an extraordinary one. In August, 
1891, I came on the Arlberg pass (1,800 m.) across a 
sanguinea colony that had rufas as slaves. Forel? has 
long since recorded some very interesting instances of 
natural, abnormally mixed colonies of these marauding 
ants in Switzerland, namely a sanguinea nest with 
F. pratensis as slaves, and another with rufa. The 
sanguineas therefore display the same peculiar uni- 
.versality and the same gift of suitable adaptation in 
their enslaving habits both in Holland, and in Tyrol 
and Switzerland; those qualities are due to that 
specific nature of their sensitive cognition and appetite, 
which we call “instinct.” 

The above observations on these “natural,” mixed 
colonies have made it plain enough, why the san- 
guineas accept the worker pupae of different, alien 


1) “S&tudes myrmécologiques en 1875,” p. 25 (57) and en 1886, 
p. 9 (189). 


Wars and Slavery in the Animal Kingdom. 79 


Formica species and rear them as slaves even when 
they are given to them by man. Forel’ and myseif 
have made various experiments on this point 
with artificial observation nests kept in a room and 
with nests found in free nature. It may suffice to 
mention one of them. In the summer of 1895, several 
times in succession I took a large bag of worker 
cocoons from a huge ant hill of F. rufa and emptied it 
in the neighborhood of a densely populated sanguinea 
nest,? which had but a few fuscas as slaves. In a few 
minutes the sanguine ants had put to flight the thousands 
of rufas contained in the bag with the cocoons and parts 
of the nest, had snatched the cocoons from the mouths 
of the fleeing rufas and began to ransack whatever I 
had brought of the hostile nest. For hours after, hun- 
dreds of these white “ant-eggs” were seen wandering 
from the plundered nest to the den of the robbers and 
mysteriously disappearing therein. By far the greater 
number of the rufa cocoons were reared by the san- 
guineas. This artificially mixed colony numbered, in 
1896, about 5,000 sanguineas and 8,000 rufas. The 
latter were generally busy building on the surface of 
the nest and had soon given it the appearance of a 
true rufa nest. At the least disturbance, however, 
thousands of light-red sanguineas would dart out from 
the interior to defend their common home; and thus 
the supposed rufa nest was turned into a sanguinea 
nest as if by magic. Because ants know no other home 
than that in which they have developed from the 
cocoon, these rufas, although they are in the majority, 


1) “Fourmis de la Suisse,” p. 258 ff. 
2) Colony No. 39 of the statistical map. 


80 Chapter II. 


will faithfully serve their ravishers and natural ene- 
mies, without “reflecting” on how they happened to get 
into this unusual society. 

I have had in my room, for the last twelve years, 
an artificial nest of F. sanguinea (see p. 23). This 
colony adopted as slaves the workers of all the Formica 
species to be found in Holland, viz., F. fusca, rufi- 
barbis, rufa and pratensis. The “slaves” have devel- 
oped from cocoons, which I had put into the nest 
during recent years. In free nature the same Formica 
species are found as slaves in the sanguinea nests, 
but only one or two of them at a time; in this nest, 
however, they were all united to form one colony 
under the suzerainty of Formica sanguinea. 

Yet, this “sugerainty” and “slavery” in the mixed 
colonies of ants is altogether different from what the 
same terms imply when applied to human society. 
Only authors like Ludwig Buechner might be found 
guilty of confounding ideas to such a degree. There 
is perfect equality among all the workers of a mixed 
colony, no less than among all the workers of a 
simple colony. The very same “constitutional laws” 
are in force both for masters and slaves; in other 
words, the uniform “nest smell,” which adheres to all 
ants reared in the same nest, serves them to recognize 
one another as members of the same ant community, 
the differences in species being totally disregarded. 
The so-called slaves live entirely free in the nest of 
their ravishers, that is to say, they live according to 
the same innate instincts which would have formed 
their rule of conduct at home; they work for their 
ravishers, supply them with food and rear their off- 


Wars and Slavery in the Animal Kingdom. 81 


spring, as if they were in their own colony. They are 
called “slaves,” only because they are reared from 
robbed pupae, live in the nests of strangers and work 
for them. On the other hand the sanguineas are 
called “masters,” only for the reason that they have 
robbed the pupae of an alien species, from which 
their auxiliaries originate; and besides, because these 
mixed colonies contain not workers only of F. san- 
guinea, but also their males and females, whereas 
the slave species is represented only by workers. This 
is why in mixed colonies the propagation of the 
masters is ensured but not that of the slaves. 
Therefore it is downright nonsense for Buechner 
to place slavery among ants and human slavery on 
essentially the same level. By virtue of his intelli- 
gence, man possesses the power of reflecting on his 
origin and social position; he is gifted with self- 
consciousness; accordingly he considers slavery as an 
unjustified deprivation of freedom, a state of humilia- 
tion, a degradation of his human dignity. With ants 
it is different. They have neither intellect nor self- 
consciousness, and are thus incapable of pondering 
over the obscure question “whence” and “whither.” 
As auxiliary ants they follow their social instincts 
just as well as in a nest of their own species: they 
are as free and independent as any other ant on 
earth. Hence among slave ants there are no run- 
aways, no revolutionists, no conspirators, no anarch- 
ists. He, who seriously points to the complete social- 
‘ ism and communism of mixed ant colonies as models 
for human socialism and political economy, is sadly 


in need of a nerve specialist. 
6 


82 Chapter II. 


On the other hand, in opposition to those animal 
psychologists who rank the “mental faculties” of the 
higher vertebrates incomparably above those of ants, 
it is necessary to emphasize the fact, that no associa- 
tion of apes or other mammals can compare, as to 
psychic faculties, with the mixed colonies of ants, 
especially with the sanguinea colonies. The rearing 
of the offspring of closely allied species as useful 
members of their own society is an arrangement never 
found with apes. The wars, therefore, and military 
expeditions of ants addicted to this practice rank 
_ much higher than the wars of baboons and other apes. 
True, slavery among ants is based only on instinct, 
not on intelligence. But anything higher than instinct 
is not found in the societies of higher animals either. 
In fact, the development of their social instincts is 
rather far inferior to that of ants. 


5. Other Wars and Alliances of Ants. 


The slavemaking expeditions of the Amazons and 
sanguineas are indeed the most interesting; but by no 
means the only wars waged by ants. There are many 
other feuds and skirmishes, both between ants of dif- 
ferent species and between different colonies of the 
same species. Most of these feuds are caused by dis- 
putes about subterranean or open-air boundaries, call- 
ing for settlement “at the point of the sword.” When 
underneath a large stone there is a “compound ant 
nest,” i. e., when two or more different species have 
built their respective nests in close proximity, they are 
separated by walls of earth. No one ventures into 
the neighboring realm, and woe to him, if he does; 


Wars and Slavery in the Animal Kingdom. 83 


he is seized and put to death. As a rule, only the 
dwarfish, yellow, thieving ants (Solenopsis fugax in 
Europe and S, molesta in N. America), may pilfer in 
the neighboring nest of a larger species. The small, 
black lawn-ant (Tetramorium caespitum, in Europe 
and N. America) will occasionally do the same. These 
thievish little ants are in the nests of larger species, 
what rats and mice are in the abodes of man. 

To return to the “compound nests.” By turning 
over the stone, under which several different ant 
species live side by side in separated nests, the par- 
titions are suddenly removed, and a fierce battle 
ensues with great loss of life on both sides. The 
engagement frequently continues for a considerable 
time after the stone has been replaced. Only after 
the boundary-lines have been perfectly restored, is 
there again peace between the neighboring states. 
Sometimes on such occasions it becomes evident that 
one colony is numerically far superior to the other. 
In this case we notice that the weaker is simply driven 
out of his nest, which is then entirely or in part occu- 
pied by the victor. Thus the war ends with the 
“territorial expansion” of the more powerful state. 

Above-ground disputes between neighboring ant 
colonies, not infrequently lead to fights, which last 
for weeks or even months, interrupted by longer or 
shorter periods of “armistice.” These battles are most 
obstinate and bloody between different colonies of the 
. small, black lawn-ants (Tetramorium caespitum). 
This species is found everywhere, frequently with 
hundreds of thousands of earth-nests within a square 
mile. On July 8, 1886, I saw a regular battlefield on 


84 Chapter II. 


-a sandy road near Exaten. It fairly swarmed with 
fighting lawn-ants. The combatants numbered thou- 
sands and they covered a space of about 70 cm. by 
8 cm. So dense was the battle-array that individuals 
could scarcely be distinguished in the mass of war- 
riors. They formed irregular clusters of from 2 to 14 
individuals all clinging together with their mandibles 
and making liberal use of ‘their stings. The summer 
heat had inflamed the rancor of the two tribes, long 
living too close together. The battle probably ended 
with the expulsion or the utter extermination of one 
of the communities. 

Among men civil wars are generally the fiercest 
and bloodiest. The same may be said of the wars 
waged between different ant colonies of the same 
species. However, only the “heat oppressed brain” of 
Buechner or Brehm could detect a closer analogy 
between these phenomena. As the males of certain 
birds fight for their breeding districts, nor allow 
other families of the same species to settle there,’ so 
ant colonies are wisely compelled by the laws of nature 
to regard the district about their nest as exclusively 
their own, on which no other colonies of the same 
species are suffered to encroach. Otherwise, their 
wants being equal, their food supplies would become 
scarce. Hence arises an instinctive hatred between 
different colonies of the same species; whereas col- 
onies of different species whose mode of life and 
means of sustenance are different are admitted much 
more easily. The preservation of the species neces- 
sitates the fiercest struggles for existence between 


1) Altum, ‘Der Vogel und sein Leben,” (6th ed.), p. 128 ff. 


Wars and Slavery in the Animal Kingdom. 85 


tribes of the same species. Not inordinate greed in 
the individuals nor imperialistic tendencies in the tribe, 
but higher, natural laws are the mainsprings of these 
“civil wars” among ants. The poet indeed, may 
exclaim: “There is room on earth for all’ (Schiller) ; 
but even in the life of ants this beautiful saying is 
often correct only in theory. 

Many more accounts of wars and battles among 
ants could be mentioned; but we cannot enter upon 
them here, since our principal purpose is to call atten- 
tion to a few points of comparison between the “intel- 
ligence” of ants and that of higher animals and of 
man. It remains only to be stated that the wars of 
ants sometimes end in an “alliance,” that is to say in 
a peaceful union of the combatant tribes into one con- 
stitutional body. These alliances are usually formed 
between Formicas of the same or different species, 
but are most frequent between different colonies of 
sanguineas. From Forel’s “Fourmis de la Suisse” 
and from my own observations (see Die zusammen- 
gesetzten Nester p. 146-157) many instances might be 
selected. The chief conditions for such alliances 
between hostile ant colonies are, that the two oppo- 
nents be closely allied in species, that they be almost 
equally populous, and lastly that they be forced to 
live in close proximity and are thus unable to avoid 
each other. Under such circumstances their original 
skirmishes give way to mutual toleration and finally 
to friendly intercourse. A superficial observer, of 
such occurrences, might be led to believe that intelli- 
gent reflection had caused the animals to overcome 
their instinctive aversion. He might conclude that 


86 Chapter 11. 


ants reason thus: ‘Why this useless shedding of 
blood? Let us not exterminate each other but live in 
peace; our differences are not so great that we can- 
not come to terms!” Yet, there is not a shadow of 
proof that ants entering into an alliance reason in this 
manner. The phenomenon, which is indeed singular 
enough, can be explained more simply and naturally 
from the laws of instinctive sensation, with special 
regard to the feeler sensations. Especially with the 
Formica species, and among these, most of all with 
the highly endowed F. sanguinea, the hereditary dis- 
position of the sensitive powers of cognition and appe- 
tite is so plastic, that with parties of almost equal 
numbers fear will be stronger in such cases than 
love of combat. First, of course, by tapping one 
another with their feelers they find out that they are 
strangers, and therefore they try to avoid each other; 
but if this is impossible, the perception of mutual 
similarity will gradually prevail over their mutual dif- 
ference. In the beginning, they live together from 
necessity only, but they gradually acquire a common 
nest-smell which unites them as members of one 
colony. From this time, by taps of their feelers, they 
recognize one another as belonging to the same house- 
hold. The former opponents have united into one 
“constitutional body” which is kept together by the 
common nest-smell. Strange though this mode of 
communication may appear to us who are not pro- 
vided with antennae, it alone explains the fact other- 
wise wholly inexplicable, how the confederate colony 
thus formed will in future hold together even against 
former members of their own colony.t An example 
~~ 4) On the explanation of the nest-smell see above p. 16 ff. 


Wars and Slavery in the Animal Kingdom. 87 


of this kind is related by Forel in his “Fourmis de la 
Suisse.’ One day he brought a handful of F. pra- 
tensis to a confederate colony of sanguineas and pra- 
tensis; the pratensis of this colony, two months 
before, had been taken from the same pratensis nest 
to which the new arrivals belonged. What happened ? 
Immediately the new pratensis were fiercely attacked 
by the sanguineas, because they were recognized as 
enemies by means of the “smell at contact.”? The 
old pratensis seemed to recognize their sisters but 
faintly. They met them with suspicion and did not 
assist them, although on the other hand they did not 
take part in the fight. But soon they began to carry 
the new arrivals into the confederate nest, as if they 
belonged to it. The number of the pratensis was thus 
increased considerably, outnumbering even the san- 
guineas. Although the latter continued their hos- 
tilities against the newcomers for several days, and 
mutilated and killed several of them, it never occurred 
to the old pratensis to make common cause with their 
maltreated “sisters” against their natural enemies. 
They allowed the sanguineas to have their own way, 
until the survivors had gradually acquired the nest- 
smell of the confederacy. In the course of a week 
“peace” was restored, and the strangers were treated 
henceforth, also by the sanguineas, as inmates of the 
same colony. 

If ants had the power of rational reflection, if they 


1) Both had been taken from the pratensis nest when fully de- 
veloped and not as pupae. Otherwise this example would be out of 
place when speaking of “allied colonies.” 

2) By this term Forel expresses very well the peculiar sensation of 
ant feelers, 


88 Chapter II. 


had any idea of consanguinity, then this behavior of 
the old pratensis of that confederate colony during the 
maltreating of their sisters would be altogether inex- 
plicable. On the other hand, the instinctive nature 
of ant sensation will furnish a very satisfactory expla- 
nation of this phenomenon, which is in evident con- 
tradiction with animal intelligence. Yet, it should 
not be forgotten, that societies of apes and other 
higher animals have nothing to compare with the con- 
federacies of ants. No one has as yet observed, that 
wars carried on between different hordes of apes 
ended with a peaceful alliance between the combatants. 
This chearly shows, how wrong it is to exalt the 
societies of higher animals above those of ants in the 
matter of psychic endowments. 

Evolutionists, therefore, such as Darwin and Zieg- 
ler, are sadly mistaken when they point to the battles 
sometimes waged between hordes of apes, and adduce 
these as conclusive evidence, that the societies of 
higher animals are so closely allied to the “primitive” 
societies of man, that a little, unimportant “step” 
bridges the difference; for first they imagine a 
“primitive state’ of human society, which is depicted, 
of course, as brutal and as devoid of reason as 
possible; then, to match the picture, they exalt the 
societies of higher animals to the greatest possible 
similarity with reasonable man, and finally, from this 
twofold hypothesis they draw the conclusion that 
human society has evidently developed from the ani- 
mal societies. And this is called the “consistent, 
scientific application of the theory of evolution to 
man!” If ants were endowed with reason and risi- 


Wars and Slavery in the Animal Kingdom. 89 


bility, they would surely burst into a hearty laugh at 
these evolutionistic “steps”; for, as to the develop- 
ment of social instincts, ant colonies bear a far closer 
resemblance to the human societies than the hordes of 
apes; and yet even the intelligence of an ant would 
be sufficient to understand, that animal and human 
societies*are as far apart as heaven and earth. The 
difference between ant states and human societies is 
readily acknowledged; but the difference between 
hordes of apes and the primitive states of man cannot 
be conceded, because, forsooth, it is against the theory 
of evolution! 


CHAPTER III. 
ARCHITECTURE IN THE ANIMAL KINGDOM. 


1. A General Survey of the Building Activity of 
Animals. 


HE houses of animals are exceedingly simple and 

destined for very prosaic purposes; they merely 
serve the wants of daily life, the preservation of the 
individual and of the species. To their owners they 
are necessary helps in the struggle for existence; they 
never aim at art for art’s sake.t This clearly shows, 
that in the animal kingdom we can speak only meta- 
phorically of architecture properly so called. There 
is mere mechanical skill, but not art; and if some- 
times its productions bear a faint resemblance to works 
of human art, the aesthetic effect is never either intended 
or understood by the animal. Another essential point of 
difference between the artistic skill of animals and of 
man is in this, that with animals it is due to an innate, 
hereditary aptitude which has not first to be acquired, 
as is the case with man. At its birth the animal is 
endowed with all its artistic talents. It applies them 
without previous experience or instruction, as soon as 
demanded by its organic development and by external 
circumstances. The caterpillar of the emperor moth 
(Saturnia) begins to be an artist only, when the time 
has arrived to transform itself into a chrysalis, and to 
weave a bottle-shaped case wherein it is to undergo 


1) The buildings of the Australian Tectonarchinae are no exception 
to this rule, if we divest descriptions of them of all poetical additions. 


90 


Architecture in the Animal Kingdom. 91 


‘the change. The female of the leaf-rolling beetle 
(Rhynchites betulae) becomes an artist, only when, the 
time of depositing her eggs having arrived, she is 
forced to cut and to roll up a birch-leaf in the shape of 
a graceful funnel. The male of this beetle is no more 
endowed with this wonderful, technical skill, than the 
emperor moth in the state of imago is able to spin a 
cocoon; for the artistic talents of animals depend on 
their organico-psychic constitution. Their application 
affords no trace of reflecting reason or free choice, 
because their mode of action is predetermined to the 
most minute detail by the laws of vegetative and sen- 
sitive life. Again, animals by their very organic con- 
stitution possess all the tools necessary for exercising 
their natural talents. Bees have their honey-bags to 
gather in the pollen and daggers to ward off their 
enemies; silkworms have their spinning glands, 
beavers use their tails as trowels, and their sharp 
gnawing teeth serve them as axe and chisel in work- 
ing the wood. There is no need therefore, to invent or 
manufacture implements. Moreover, directions for 
using their natural instruments are furnished to ani- 
mals by the innervation of their bodily organs and by 
the corresponding constitution of their sensitive powers 
of cognition and appetite. Reason and free choice, 
therefore, are utterly superfluous in the exercise of 
these animal talents. Since by their organico-sensi- 
tive constitution animals are provided sufficiently with 
all the necessaries of life, they are denied the higher, 
spiritual faculties. Hence, they, who attribute reason 
to animals, show that they have but a superficial 
knowledge of animal life. 


92 Chapter ITI. 


The buildings of animals either serve to shelter 
the individual, or else they are places for breeding 
and rearing the young. To the former class belong 
the tunnels excavated in the earth by the serpulas, the 
envelopes made of various substances by the moth- 
caterpillars and the larvae of the may-flies, as also 
the different casings constructed by the larvae of 
insects, particularly of many butterfly-caterpillars 
before their metamorphosis. To the latter class 
belong the regular nest constructions of animals. The 
most primitive specimens are found with the parasitic 
Nemertine worms. In several orders of insects, 
especially among the Hymenoptera? and the beetles, 
we meet with instances of ingenious and manifold 
development of the same art. Here we find the most 
various forms of nests, and made of all kinds of 
material. Those elegant little domes of mortar, the 
wasps of the genus Eumenes have built for their 
offspring. Those breeding burrows, lined with red 
poppy blossoms, have been’ excavated by the so-called 
tose-bee (Megachile). Those graceful funnels and 
barrels of leaves have been rolled into shape for their 
young by weevils (Rhynchites, Apoderus, Attelabus). 
And that boat adorned with streamers has been spun 
by the great water-beetle (Hydrophilus piceus), as a 
receptacle for its eggs. Among fishes nest building is 
tare. We find examples of it, in the stickleback (Gas- 
terosteus aculeatus), and in other fishes provided with 
spines. On the other hand, birds are unsurpassed in 
the art of nest building as regards variety, both of 





1) See “Naturforscher,” 1886, 19th year, No. 50, p. 494. 
*) See especially J. H. Fabre, “Souvenirs Entomologiques,” who 
has described these buildings with admirable skill. 


Architecture in the Animal Kingdom. 93: 


form and of the material used by different species. 
With the mammals,! finally, nest constructions are, on 
the average, far less complicated and artistic than with 
birds and insects. 

The buildings which serve to shelter and rear the 
young, may likewise be used as permanent lodgings 
for parents and offspring. This is the case with social 
insects and many mammals. Thus the nest develops 
into a family dwelling. Only in relatively rare 
instances, do animals employ their building skill in 
providing other necessaries of life. Many spiders spin 
their webs not only as a hiding-place for themselves 
or for breeding purposes, but they also, by means of 
their spinning glands, manufacture nets wherewith 
to catch their prey. In like manner the neuropterous 
larva, which goes by the name of ant-lion, uses its 
earth-funnel both as a dwelling place and as a trap for 
catching its prey, which consists chiefly of ants or 
other insects. Among ants, however, we find the most 
varied and manifold application of natural architec- 
tural skill. 


2. The Nests of Ants. 


In the first place, ants use their architectural talents 
for building nests, in the strict sense of the word. 
Everybody is more or less familiar with ant nests, but 
few are aware of the immense variety of forms implied 
in that apparently simple term. As there is scarcely 
any material unfit for an ant nest, so it may assume 
all possible shapes and be found in the most unlikely 


1) We shall consider more in detail the buildings of the beavers in 
the subsequent pages. 


94 Chapter ITI. 


localities. Here it is the size of a thimble, there, the 
pyramids of the ancient Egyptians are like mole-hills 
in comparison, if we take into consideration the rela- 
tive size of the builders. Some are in the ground, in 
clefts of rocks, or concealed by stones, others are 
under the bark or in the wood of trees. Others again 
are in the hollow stalk of a plant, or in a gall-nut or 
in a deserted snail-shell. Now they hang high in the 
boughs of a tree, now in forests they rise as domes 
from the level of the ground. Such a nest may be 
dug, or spun; it may consist of masonry, or of cavities 
hollowed out of the earth or of the wood. Sometimes 
all these modes of operation enter into the same con- 
struction. In short, the variability as to form, style, 
or locality is almost unlimited. There is one charac- 
teristic, however, common to all ant nests, viz: the 
absence of any uniform architectonic pattern; ant 
nests are irregular systems of chambers and galleries, 
giving shelter to the ants and their offspring, and 
communicating by different openings with the outside 
world. This very irregularity of their buildings 
enables the ants to suitably adapt their nests to any 
locality and to employ any kind of material in their con- 
struction. The artificial and, as it were, mathematical 
regularity of the honey-combs of bees? is entirely 


1) N. Ludwig, in an essay, “Der Zellenbau der Honighbiene,” (in 
“Natur und Offenbarung,” 1896, 10th issue, p. 598 ff.), has offered a 
new explanation of the hexahedral form of the bee-cell and of the three 
congruent trhombs forming its pyramid-like base. In his opinion the 
peculiar form of the bee-cell is due only to the construction of the 
wax combs, each cell being built only in connection with other cells. 
For the bees are actuated by the impulse of combining round cell-walls 
bordering on one another, into one single wall and to reduce their 
thickness by gnawing off both sides as much as possible without peril 
to their necessary strength. Hence, the flat walls of the form described 


Architecture in the Animal Kingdom. 95 


wanting. This difference is very important for com- 
parative psychology. As in the social life of ants 
the individual independence of the single workers 
attains a higher degree than amongst bees, so in their 
architecture the same phenomenon may be observed. 
Instead of constant uniformity we find great variety; 
instead of the monotonous “automatism” of innate 
instinct, we meet with a quasi-intelligent arbitrariness 
in the exercise of their sensitive cognition and appe- 
tite. Scarcely anywhere else is the wonderful plas- 
ticity of animal instinct manifested so clearly and con- 
vincingly as in the architecture of ants. This was the 
reason why ants seemed to furnish such numerous 
proofs of the “individual intelligence” of animals; 
and, in fact, im the architecture of ants, if anywhere 
in the whole animal kingdom, we find a striking 
resemblance to human intelligence. 

In order to give a full psychological description of 
ant architecture, we shall first, compare the nests of 
different species, secondly, we shall consider the differ- 
ences in the buildings of one and the same species; in 
the third place, we shall examine the methods by which 
ants of the same colony co-operate in building their 
nest, and finally, we shall investigate the various pur- 
poses to which ants apply their architectural talents. 
However, it would require a volume of considerable 
bulax to treat these points exhaustively. Therefore, we 


above. The consequently elongated, prismatic form of bee-cells would 
be merely the result of the extreme uniformity in the working of the 
bees engaged in building the combs. Single cells are built by the same 
bees always in a cylindrical form with hemispherical top and bottom. 
This form is to be regarded, according to Ludwig, as the proper, 
primitive type of bee-cells, which is found with bumble-bees and other 
allied Hymenoptera. 


96 Chapter ITI. 


must confine our discussion to its narrowest possible 
Simits, touching chiefly on such features as are of 
special interest for comparative psychology. 

Great as is the variety of ant nests, still, every 
species has its peculiar architecture, differing more or 
less from that of any other species. Many ants, e. g., 
our small, blackish garden ants (Lasius niger) and the 
small, yellow meadow ants (Lasius flavus)* work 
almost exclusively in earth. Their nests are dug in 
the ground, but above the subterranean nest they raise 
smaller or larger domes of earth, the stalks and blades 
of grass, that grow on the spot, serving as natural 
pillars and beams. Other species, again, e. g., our well- 
known hill ants (Formica rufa)? build so-called “ant- 
hills,’ the popular type of ant nests in our northern 
hemisphere. These ant hills may be termed mixed 
buildings, an under-ground earth nest being combined 
above ground with a dome consisting of earth, pine- 
needles, scraps of dry leaves and stalks, and other parts 
of plants. The different ant species which build such 
ant hills follow systems and styles peculiar to each. 
Thus any one with a little practice is able to determine 
at once the species of the builders. F. rufa builds 
differently from pratensis, pratensis from e-xsecta, 
exsecta from sanguinea. The universal tool which, 
like the human hand, is fit for and skilled in a 





1) Both are found also in N. America; the most common yellow 
ant there is L. aphidicola Walsh; L. niger is represented by its very 
common N. American race L. americanus Em, 

2) The N. American species of the rufa group are very numerous; 
among them the nests of F. exsectoides For. and obscuripes For. are 
most like the European ant hills of F. rufa.—An interesting descrip- 
tion of N. American nests is contained in a paper of Father H. Mucker- 
mann, S. J., entitled ‘‘The structure of the nests of some North Ameri- 
can species of Formica.” (“‘Psyche,” June, 1902.) 


_ Architecture in the Animal Kingdom. 97 


variety of performances is found in the jaws (mandi- 
bles) of the ants. Of course, in digging burrows in 
the earth and in constructing earth-works they are also 
assisted by their fore-legs, which help partly to scrape 
up the sand and partly to hold down and fasten pellets 
of earth. In closely allied species the shape of these 
instruments, and especially that of the all-important 
toothed inner edge (cutting edge) of the mandible, 
is as a rule so similar* that the specific differences in 
architectural style can be accounted for only by the 





Fig. 2. Fig. 3. 
Left mandible of Formica rufa. Right mandible of F. sanguinea. 
orker.) (Worker.) 


instinctive preferment of a particular style on the part 
of different ant species. In the case of ants, therefore, 
it will never do to resort to the mechanical automatism 
of animal activities, and to explain the differences of 
instincts merely by differences of bodily organs. The 
decisive factpr is the psychic variety of instinctive dis- 
positions. By them the bodily organs, in themselves 
indifferent, are directed in their various modes of 
operation. 

It is true, to a certain extent, that the nature of 


1) See the subjoined cuts. Both are drawn with the Zeiss’ 
microscope, syst. A,, and Abbe’s Camera lucida. 
7 





98 Chapter III. 


the exterior organs of ants will also decide the nature 
of their architecture. Thus e. g., the large Camponotus 
ligniperdus (horse ants) and their allied species pos- 
sess larger workers, whose huge head and strong 
mandibles enable them to cut galleries in the wood of 
decayed or even of sound trees. And therefore these 
species are remarkable for wood nests. Others again, 
among them the jet black Lasius fuliginosus as the 
only one of this kind among the emmets of northern 
Europe, build paper nests by gnawing wood-fibre and 
gluing it together with the sticky product of their 
salivary glands. They thus produce a coarse, brown 
papiermaché, in which they establish their nests. Far 
more perfect are the paper nests made by several 
foreign ants, especially in South America, Madagas- 
car and East India. They resemble irregular, brown 
or grey-colored wasp-nests, suspended from or fast- 
ened between branches of trees. Rev. A. Schupp, 
S. J., sent me from Porto Alegre (South Brazil) 
several paper nests of Cremastogaster sulcata, one of 
which on arriving in Holland still contained several 
thousands of live inhabitarits. Similar nests of Cre- 
mastogaster Schenki in Madagascar are reported by 
Sikora to be sometimes of such size as to accommodate 
a full-grown man. From these paper-nests we must 
distinguish nests which are spun and do not consist of 
a paper-like material but of a texture like cobwebs. 
Such webs are constructed, according to Wroughton’s 
observations,| by an East Indian ant Polyrhachis 
spinigera for lining her earth-burrows. Other Indian 
and Australian ants of the genera Oecophylla and Poly- 


1) “Our Ants,” part I, p. 25 (“Journal of the Bombay Nat. Hist. 
Soc.,”’ 1892). - % 


Architecture in the Animal Kingdom. 99 


rhachis build their nests on trees by sewing together 
clusters of leaves. It is now finally ascertained (see 
p. 128-129), whence the Oecophylla procure the 
material for their threads. They do not secrete it 
from their mandibular glands, as was hitherto sup- 
posed,’ but they use their young larvae as “spinning- 
wheels,’ moving them to and fro between the edges of 
the leaves. This strange fact observed already by 
Ridley and Holland? has recently been confirmed by 
other scientists. 

Another class of ant nests, likewise found in the 
tropics exclusively, are the natural cavities in the 
stems, thorns and bladder-shaped swellings of the so- 
called “ant-lodging plants,’ which invite ants to take 
possession of their well furnished lodgings. Several 
of these plants, as the American Imbauba (Cecropia 
adenopus) offer to the ants, besides the lodging, also 
an agreeable food in the form of special honey-bearing 
nectaries. In return for their kindness the “ant-lodg- 
ing plants” are afforded by their valiant lodgers effec- 
tive protection against leaf-cutting ants and other 
herbivorous insects. This mutual relationship of the 
ants with the plants in question is called Symbiosis 
(consociation). It bears, in fact, some similarity to the 
associations existing between animals of different 


1) E. H. Aitken, “Red Ants Nests” (“Journal of the Bombay Nat. 
Hist. Soc.,’”? 1890, Vol. V, n. 4, p. 422), also Forel, “Die Nester der 
Ameisen” (Zuerich, 1892), p. 19. 

2) E. E. Green, “On the Habits of Oecophylla smaragdina F.” 
(“Proceedings Entomol. Soc. of London,” 1896, p, IX.) 

8) Fritz Mueller, “Die Imbauba und ihre Beschuetzer’’ (‘‘Kosmos,” 
VIII, 109), and A. F. W. Schimper, “Die Wechselbeziehungen zwischen 
Pflanzen und Ameisen im tropischen Amerika’ (Jena, 1888). Also 
H. v. Jhering, “Die Ameisen-von Rio Grande, do Sul,” in “Berl. 
Entom. Zeitschr.,” 1894, 3d issue, p. 354 and 364 ff. 


100 Chapter ITI. 


species, e. g., between ants and their guests, such as 
the club-bearing beetles (Claviger), the tufted beetles 
(Lomechusa, Atemeles), etc. But even those cavities 
of plants, which are properly not meant to receive and 
to lodge ants, are often occupied by them, especially 
_ in the luxuriant vegetation of tropical South America. 
Aug. Forel in the winter of 1895 and 1896 visited 
the savannas of Columbia, where he found that the 
nests of by far the most of the species, belonging to 
eight different genera, were built indry stalks of grass.* 
This led him to think that in the prairies and forests of 
tropical America the nests in stalks and in hollow parts 
of plants were the typical form of antnests, correspond- 
ing to the climate of that country, whereas in our 
zones the usual type is the earth nest or else the hill 
made of earth and parts of plants. 

This cursory comparison of the various forms? of 
nests met with in different ant species, shows clearly 
enough, that their character is conditioned by the 
peculiar shape of the bodily organs of the builders , but 
far less than is the case with most of the other artistic 
instincts in insects and other animals. The form of 
the mandibles, the presence of salivary glands with 
gluey secretions or of real spinning glands, indicate 
only the general outline of the architectural style 
preferred by their owners. Only the different instinc- 
tive dispositions of the builders determine more 
exactly the specific differences of their nest forms. 


1) “Quelques particularités de l’habitat des fourmis de 1’Amérique 
tropicale’” (Extr. des Ann, de la Soc. Entom. Belg., XL [1896], 167 ss.) 
and “Zur Fauna und Lebensweise der Ameisen im columbischen 
Urwald,” in “Mitteil. der Schweiz. Entomol. Gesellsch.,” IX, 9th issue, 

2) Forel, “Die Nester der Ameisen,” Zuerich, 1892. 


Architecture in the Animal Kingdom. 101 


Within the limits of these instinctive, hereditary dis- 
positions there is plenty of room for the worker ants 
to give full play to their individual powers of sensitive 
cognition and appetite. This is why certain ant spe- 
cies, particularly those skilled in earth work or wood 
work, often take possession of some spot so suitable 
for their dwelling as to be practically half-finished. 
This they fit up in a becoming manner as a home for 
their colony. Such attractive spots are e. g. on the 
heaths of northern Europe rotten tree-stumps, in which 
for years numbers of bark-beetles and their larvae or 
other wood-boring insects were kind enough to prepare 
comfortable quarters by carving out an extensive sys- 
tem of galleries and chambers. It only remains for 
the ants to take possession of the lodgings, devour 
the former inhabitants, if there be any left, clean the 
- apartments, close up the superfluous entrances with 
earth or rotten wood, and with the same materials 
construct, if need be, a few partitions for separate 
chambers. If, on occupying the residence, they should 
perchance discover that part of the stump is already 
inhabited by another ant colony, the latter are killed 
or turned out without much ado. If they should fail 
in this, they make peace and live on good terms with 
the rivals, especially if the latter are equal to them in 
fighting strength, but differ greatly in size and means 
of defense. The nests are separated by partitions, and 
the stump is henceforth inhabited by a “compound 
nest.” 

Such “stolen nests” iorm a large category of ant 
nests, particularly in places where there are many 
stones; for almost all ant species that dwell in earth 


102 Chapter ITI. 


nests, have a predilection for building under stones. 
This saves them a great deal of work and gives the 
whole building greater firmness, and, moreover, in 
. such a nest the heat of the sun more easily penetrates 
to the interior. In heaths also, where stones are rather 
rare, instances of such stolen nests may occur. A nest 
of F. sanguinea,’ on which in 1894 I had placed a clod 
of heath serving it henceforth as roof, had in 1895 
passed into the possession of a colony of Lasius niger ; 
in the years 1896-98, it was again inhabited by F. san- 
guinea. A short time ago, in the same region near 
Exaten, I found a rather extensive earth hill supported 
in the centre by a bunch of heather and inhabited by 
a large colony of F. rufibarbis. The ants had collected 
on the surface a small heap of dry heather-leaves, as 
they generally do there. The earth hill itself, how- 
ever, judging by its architecture, was an old nest of 
L. niger, which subsequently had been occupied by 
the rufibarbis. History is silent as to whether the orig- 
inal builders of the nest had quitted it before the time 
_of the foreign invasion, or whether they were com- 
pelled by force to evacuate it. 

F, sanguinea are a restless people. They frequently 
desert their nests, which are then taken possession of 
by other, smaller species (especially Tetramorium 
caespitum, L. niger and alienus). I have noted a num- 
ber of such instances in my records of the last few 
years; it may suffice to mention one of the most 
remarkable. A large colony of F. sanguinea (No. 72 
of my stat. map), in 1895 and 1896, had inhabited a 
group of three nests, distant from one another 3 and 


1) Colony No. 155 of the statistical map. 


Architecture in the Animal Kingdom. 103 


ym. respectively. Since the spring of 1897, however, 
they had emigrated and I did not find them again. 
In 1898 the northernmost of these nests was empty, 
the central one was occupied by L. niger, and the 
southernmost by Tetramorium caespitum. This was 
the state of affairs until July, 1898. When I returned 
on July 14, I found that the sanguineas of colony No. 
72, which I could easily recognize by the size of their 
workers, had returned to the southern nest which con- 
sisted of two little heaps in close proximity. From one 
of these the Tetramorium had already been completely 
driven out; in the other they still occupied a retired 
corner and were surrounded by the sanguinea nest. 
Therefore ant nests also have their fates. 

Ant nests are stolen not only by ants of different 
species, but sometimes by those of different colonies 
belonging to the same species. Many instances of this 
kind are furnished by the sanguinea colonies in the 
neighborhood of Exaten; but for brevity’s sake, it 
must suffice merely to mention the fact. 


3. The Nests of the Sanguine Slavemakers. 


The great plasticity and power of adaptation to 
given circumstances in the nest-building instincts of 
ants is seen to the greatest advantage by an examina- 
tion of the nests of the sanguine slavemakers (F. san- 
guinea). With members of one and the same species 
possessing the same specific, natural constitution, there 
is such a variety of nest construction, that there is no 
trace of that “automatism” of instinct, which postu- 
lates a completely uniform and monotonous manifes- 


104 Chapter III. 


tation of the hereditary instinctive activities. As 
these ants, in the parlance of modern animal psychol- 
ogy, possess a high degree of “individual intelligence,” 
because under the influence of their sense-perceptions 
and sense-experiences they, are able to adapt their 
innate, instinctive dispositions and aptitudes to any kind 
of circumstances, so they manifest great adaptibility in 
the building of their nests. I have drawn up statistics 
of the sanguinea colonies in the neighborhood of Exa- 
ten, which show that to my knowledge there are in 
this region about 2,000 nests of this ant species, 
embracing 410 colonies. The architectural style of the 
nests varies greatly. By far the majority of them 
are underground, built either below the bare surface 
or under a shrub of heather, beneath a loose clod, 
under a stone or at the foot of a tree. In connection 
with this underground earth-nest there is generally 
on the surface a greater or smaller heap of dry leaves 
collected from the heather shrubs. This heap, together 
with the earth carried out of the interior galleries and 
the twigs of the shrubs supporting the whole construc- 
tion, forms a sort of protective dome. With large nests 
this hill sometimes has a circumference of several 
meters and a height of several decimeters (e. g., in 
colonies Nos. 208, 216, 118); but often it is rather 
insignificant and sometimes it is altogether wanting. 
Besides these simple or mixed earth-nests, F. san- 
guinea builds also in rotten stumps of fir-trees or oaks, 
now under the loose bark, now in the wood, now in the 
roots. Sometimes the old stump is surrounded by 
earth-galleries, and sometimes the whole nest is con- 
fined to the stump itself. The nest of one of our 


Architecture in the Animal Kingdom. 105 


sanguinea colonies here (No. 112 of stat. map) is 
built in a lofty and sturdy oak, in the mould and the 
clefts of a hollow in the tree, almost a yard above the 
ground. These observations show that the sanguine 
ants are able to choose the most peculiar places for 
their nest and to adapt its construction to any local 
circumstances. This is confirmed by the following 
observation. In the immediate neighborhood of Exa- 
ten the earth-nests prevail, while two kilometers further 
or near the village of Grathem, the greater number of 
nests are built in rotten fir-stumps, because in that 
region the ground is more turfy and thus renders the 
construction of earth-works rather laborious. 

Not less variable than the style is the number of 
nests constituting a colony of F. sanguinea. Among 
the above mentioned 410 colonies which are known 
to me in this region, there are but a few that have only 
one nest. These are mostly weak tribes which, on 
account of the scarcity of members, feel no need of 
other nests. In some cases, however, even a very 
strong colony has only a single nest built in a specially 
convenient place, generally at the foot of a fir 
(e. g., col. 208 and 216). In such cases the concen- 
tration of the building is of greater advantage than 
its division into a number of different nests. 
However, by far the majority of the sanguinea colo- 
nies have several nests, averaging from two to eight, 
either close together or farther off from another, some- 
times inhabited simultaneously, sometimes by turns. 
The distance between the nests of one colony is mostly 
only from 4 to 4 m., but sometimes from Io to 20 m, 
or more. The latter is particularly the case with their 


106 Chapter ITI. 


summer and winter residences, or rather with the 
abodes for winter and for spring; for, many but by 
no means all sanguinea colonies of this country have 
special winter-quarters established in thickets under 
the roots of trees or stumps and affording deep and 
warm recesses for the cold season. The spring resi- 
dence, however, which again often consists of several 
single nests, is generally built near the edge of the 
thicket. Here, on the first warm days of March and 
April, the ants can always be observed changing their 
quarters and moving over with the whole family to 
the spring residence. In September or at least in the 
beginning of October they change again, moving in 
the opposite direction. 

If it should become very hot and dry in summer, 
the colonies move to their winter quarters during the 
dog-days, thus converting the winter nest into a mid- 
summer nest. When at the end of August, 1898, I 
had returned after several weeks of absence, I found 
that during the exceptionally hot days of August most 
of the sanguinea colonies of this region had aban- 
doned their spring nest. What had become of them? 
As I was well acquainted with the winter nests of many 
colonies by the means of the statistical map I had 
drawn up in the preceding years, it occurred to me to 
look there in search of them. The result was rather 
striking: all of the emigrated colonies possessing 
winter quarters of their own, had already occupied 
them! This was such a regular occurrence, that, when 


1) The country about Exaten consists in its uppermost layer of 
light sand, which at once loses all its moisture in places exposed for 
some time to the scorching rays of the sun. This condition of the soil 
is surely essential in explaining the facts just mentioned. 


Architecture in the Animal Kingdom. 107 


I found the spring nest deserted, I had only to look 
where a given colony had passed the winter; there 
they would be found under the clods of their nest. 
Strange, you might say, to use one’s winter quarters 
as a summer resort! Yet, on considering the shelter 
afforded by a winter nest, situated in the brushwood 
or at the roots of a shady tree, we may easily under- 
stand that the very same place can protect ants both 
against the cold of winter and against the scorching 
heat of summer. In this winter and midsummer nest 
the sanguineas generally stay during autumn and pre- 
pare for their hibernation. Whilst the hill ants (F. 
rufa and pratensis) are still busy building their hills 
and setting out on expeditions to visit aphides, whilst 
the small, black lawn ants (Tetramorium caespitum), 
the red stinging ants (Myrmica rubra) and the small, 
brown garden ants (Lasius niger) are still in full 
activity around their nests and are accompanying their 
winged sexes on the way to their nuptial flight, there 
prevails in the nests of sanguinea a quiet in striking 
contrast with the feverish activity of the inhabitants 
during the former months. The really active season 
for F. sanguinea is from the middle of March to the 
middle of August. Therefore their spring nests also 
may be called working nests, whereas their winter 
nests, which serve for midsummer and autumn, may 
be called nests of repose. 

Besides these periodical changes of residence, F. san- 
guinea will also incidentally move to another nest with- 
' in the temperate or hot season. This moving is caused 
by special conditions of the weather. If, on account 
of long aridity and heat, the sanguine ants begin 


108 Chapter ITT. 


- 


to feel uncomfortable in their residence on the southern 
edge of a fir-plantation, they emigrate, bag and bag- 
gage, to the more shady side on the north. Toward the 
end of May, 1896, I observed several colonies chang- 
ing their nests for this reason. If the weather changes 
and continues to be chilly and rainy, they bundle up 
once more and go back to the old home. A similar 
instance, I noticed on June 20, 1896. There had been 
heavy showers for several days back and many san- 
guinea colonies that hitherto had lived in earth-nests 
were moving into old oak stumps; for these afforded 
them a better shelter against the penetrating rains. 
Those who designate as “intelligence,” every suitable 
change in instinctive activities, caused by sense percep- 
tions and sense experiences in animals, can hardly 
escape from crediting ants with rather a considerable 
degree of animal intelligence; for even in the highest 
mammals we hardly find a higher degree of “psychic 
plasticity,’ than is manifested in the above examples 
by the sanguine slavemakers. However, we have 
already shown in a former essay (Instinct and Intelli- 
gence in the Animal Kingdom), that it is entirely 
wrong to apply the term “intelligence” in this man- 
ner; for the phenomena in question can be fully 
accounted for by instinctive sensation and, therefore, 
they do not supply the least evidence in favor of intel- 
ligent, mental activity of the animal. Man, of course, 
in observing such phenomena, can attribute to the ants 
the following reasoning: “For the last few days it has 
been raining a great deal. We and our children have 
become dripping wet. Now, we do not want to get 
wet again; therefore, we must move to another dwell- 


Architecture in the Animal Kingdom. 109 


ing, where the rain can not enter; those old oak 
stumps, however, are just the thing; therefore we 
move to that place.”—We, however, maintain: With- 
out admitting animal intelligence the whole affair is 
explained much better from the instinctive association 
of sense representations. The ants do not like the 
old place any more on account of the disagreeable 
experiences undergone there, therefore they look for 
another. That under these circumstances, just those 
dry oak-stumps appear to them to be so very inviting, 
follows from the suitable disposition of the sensitive 
cognition and appetite. That ants in such cases are - 
intellectually conscious of the suitableness of this 
change of nests, is an unwarranted assumption to which 
we reply: quod gratis asseritur, gratis negatur. In 
other words: we are not allowed arbitrarily to attrib- 
ute a human course of reasoning to animals in the 
sense of “popular” psychology. Such men as L. 
Buechner may find a proof of the “high intelligence” 
of ants in the fact that, e. g., in low-lands Leptothoraxr 
acervorum resides under barks of trees, but in the 
Alps under stones.t Although we consider the power 
of adaptation manifested by the sanguineas in their 
nest-building instincts far more deserving of admira- 
tion, yet we are far from regarding even this power as 
an instance of animal intelligence, but, rather, of animal 
instinct, the various activities of which depend neither 


1) Buechner, “Geistesleben der Thiere,” p. 73. In this book the 
author calls Leptothorax acervorum erroneously Lasius acervorum. 
Romanes in his book, “Die Geistige Entwicklung im Thierreich” 
(Leipzig, 1885), p. 268, was surely referring to the same passage of 
Buechner, because he still more erroneously calls that ant Lasius 
acerborum. 


110 Chapter III. 


on mechanical automatism nor on individual reflec- 
tion of the animal, but on the suitable disposition of 
its sensitive cognition and appetite. 

True, the plasticity of the building instinct is 
greater with the sanguineas than with their allied 
species; yet even the latter sometimes perform actions 
that go to prove clearly, that the nest-building instinct 
in ants is not blind mechanism, but is suitably modified 
by their sensitive cognition. I observed a striking 
instance of this kind in the summer of 1898 at Lipp- 
springe in Westphalia. In a growth of young fir-trees 
near the so-called Fisherman’s hut there lay a small 
heap of old pieces of tar-paper. This treasure had 
been discovered by some Formica truncicola N yl., which 
had their nest at a distance of 64 m. in a fir-plantation 
on the other side of a broad, sandy road. Their nest 
was a normal truncicola nest, a hill of fir-needles and 
earth, built around a fir sapling. Now, the ants were 
better pleased with the newly discovered place under 
the tar-paper, than with their original nest, therefore 
they moved over, bag and baggage; and the moving 
lasted several weeks. The tar-paper afforded them in 
a far higher degree the advantages usually derived 
from their surface domes called ant-hills, for under 
the layer of tar-paper warmed by the rays of the sun 
_ there was a uniformly higher temperature and, at the 
same time, an effectual shelter against the rain. There- 
fore they established their nest under the tar-paper 
without surmounting it by a hill. When I returned 
to Lippspringe at the end of May, 1899, the truncicola 
nest was still under the tar-paper, no indications of 
any building being visible above ground. Their former 


Architecture in the Animal Kingdom. 111 


hill in the forest, however, was in ruins, a sign that it 
had been completely abandoned by its builders. 

Neither experience, nor instruction, nor “profound 
thought” could have taught the ants, that tar-paper 
possesses in a high degree the qualities which allowed 
them to dispense with an ant-hill; for it was a material 
which ants are not wont to meet with. It is not, there- 
fore, “intelligence,” but the instinct of ants that rightly 
accounts for a change of nests seemingly so wise. The 
first ants that happened to find the paper were de- 
lighted with the place; their senses were impressed 
with its comfort and security. This led them to bring 
over some of their nest-mates, and since these also 
were pleased, the whole colony finally emigrated and 
settled at the new place. The sensitive powers of 
cognition and appetite also explain very easily, why 
the ants did not raise a hill above the tar-paper. Under 
the artificial roof they felt safe and warm enough 
without a superstructure, therefore they saw no need 
of any additional construction. Other ants, too, often 
omit to build above ground, when they meet a 
stone, that furnishes sufficient warmth and protection 
for the underground parts of the nest. Even of F. 
rufa, which is the most typical “hill-ant,” I found a 
series of nests under stones, near Goebelsmuehl, 
(Luxemburg) in July, 1904. 

As ants, and in particular the sanguine slave- 
makers, are able to adapt their skill in building nests 
_ to the most varied localities and conditions of season, 
so the same instinct manifests great plasticity with 
regard to sudden emergencies, e. g., against the attacks 
of their various enemies. A weak colony of F. san- 


112 Chapter III. 


guinea selects for its nest a more hidden place than a 
stronger one does. This is especially the case, if there 
are hostile ant species in the neighborhood, liable to 
pay them a visit at any time. Thus colony 166 of my 
statistical map of Exaten, being harassed by neigh- 
boring pratensis, had at last hidden itself so well in 
the earth, that I was obliged to spend a considerable 
time before finding it. I myself have often enough 
been vexed to see that colonies of the same sanguinea, 
on being disturbed even by the hand of man, emigrate 
and seek elsewhere a new place of settlement. Some- 
times even a clod of heather placed on their nests will 
cause weaker colonies to emigrate. This is all the 
more strange, since our sanguinea is very fond of 
using such clods as a roofing for her nests. Other 
colonies, indeed, did not emigrate on account of my 
repeated visits, but tried to retire farther into the 
interior. A striking instance of this kind was offered 
by colony No. 36, which was of moderate dimensions. 
Originally the exterior of the nest looked like any 
common earth-nest of the same numerical strength. 
The clod which I had placed on it was used as a roof 
and was covered by a heap of dry heather leaves. I 
often visited the nest and each time I lifted the clod 
to have a view of the interior of the building. In con- 
sequence the ants blocked up the former entrances; 
they dug new ones at a greater distance from the 
nest, and came to the surface much more rarely. Even 
the small heap of materials gathered from plants 
gradually disappeared from the top of their nest. 
After a time it was blown or washed away, nor did 
the ants renew it. It was evidently instinct that led 


Architecture in the Animal Kingdom. 113 


them to withdraw from these repeated disturbances by 
closing up and concealing their nest. 

Professor Aug. Forel’ relates that he had brought 
home from Algeria a colony of Myrmecocystus altis- 
quamis and placed it in his garden at Zuerich; but 
owing to the trouble caused them by the small ants 
Lasius niger and Tetramorium caespitum they grad- 
ually modified their usual manner of nest construction. 
Under normal circumstances this Myrmecocystus 
species has wide open nest entrances; in this case, 
however, they were contracted to afford greater pro- 
tection against the thievish visitors, and finally they 
were almost entirely closed up. This instance is sim- 
ilar to the one recorded above of colony 36 of F. 
sanguinea, and is psychologically to be explained in 
. the same way. The repeated disagreeable experiences 
caused to the ants by the troublesome strangers 
induced the Myrmecocysti, contrary to their former 
habits, to close up and to conceal their nest. As 
Forel points out, these facts afford irrefutable evidence 
of the great plasticity of ant instinct. For, this 
instinct is not merely a nervous mechanism forced to 
operate along uniform lines; it includes sensitive cog- 
nition and appetite, which are not only of an organic 
but also of a psychic nature. Thus animals are 
enabled, by new sense perceptions and experiences, 
to adapt their wonted mode of action to the require- 
ments of circumstances. This does not, however, 
_ compel us in the least to attribute to animals a power 
of cognition essentially the same as human intelligence ; 


1) “Les Formicides de la Province d’Oran” (Lausanne, 1894), p. 
8; see also “‘Apergu de Psychologie comparée,” p. 24, by the same 
author. 


114 Chapter III. 


in fact, we cannot even do so, if we wish at all to 
proceed scientifically. Popular psychology may, 
indeed, perceive a “spiritual power of reflection” in 
those activities of animals, in which sense experiences 
enter as additional factors. And of course, this 
uncritical procedure forces them, in consequence, to 
ascribe to ants at least the same degree of “individual 
intelligence” as to the highest mammals; for, with 
the latter great plasticity of instinct is rarer than with 
ants. These conclusions, which are declared absurd 
by modern evolutionists themselves, clearly prove the 
untenability and’ self-contradiction of modern animal 
psychology. 


4. How do Ants Build Their Nests? 


In spite of its irregularity every ant nest is always 
a unit, consisting of one or more chambers, galleries 
and entrances, by which ants communicate with the 
outside world. And now we are confronted with the 
question: How do the members of a colony co-oper- 
ate in building their nest? 

Almost a hundred years ago Peter Huber? atten- 
tively observed the ants building their nests, and 
described in a masterly manner the skill and assiduity 
of these small animals. Any one can satisfy himself 
of the correctness of these observations by watching 
wood ants (F. rufa) building their hills on some 
sunny day in spring, or by looking at the small, black 
garden ants (Lasius niger) constructing their earth- 
nests during some warm spring shower. 


1) “Recherches sur les moeurs des fourmis indigénes” (1810). 
Nouvelle édition, 1861. Chap. I. : 


Architecture in the Animal Kingdom. 115 


In building nests the single workers co-operate 
differently in different species; nowhere, however, do 
they co-operate with the regularity of a machine or 
according to a rigid pattern, but each ant with evident 
liberty follows her own impulse and her own plan. It 
is above all the instinct of imitation, which neverthe- 
less causes a uniform result, a nest consisting of com- 
municating galleries and chambers. As a rule the 
most zealous and skillful worker is imitated most; 
her zeal is catching, so that she directs the activity of 
the others into the same channel. This mode of 
co-operation effected by means of the instinct of imi- 
tation is predominant in hill ants (F. rufa) and the 
small, blackish garden ants (Lastus niger). F. fusca, 
however, the greyish-black slave ant or negro ant, 
(Huber’s fourmi noir-cendrée) belongs to those 
species, in which the independence of the several 
workers in building is especially remarkable. The 
same is observed with the closely allied F. rufibarbis 
(Huber’s fourmi mineuse). I.often witnessed in 
these two species, how the pellets of earth, which one 
worker had just put down for building a wall in a 
certain place, were carried away by another one in 
order to apply them elsewhere in a manner more suit- 
able to her own taste. To observers, who know the 
habits and doings of ants but superficially, and are 
wont to transfer their own thoughts into the brain of 
the animal, such occurrences may appear as though 

an ant wished “purposely to correct” the work of 
~ others. And, as a matter of fact, one of P. Huber’s 
observations! bearing on this head has been actually 


1) Lic., p. 48. 


116 Chapter III. 


thus interpreted by writers on popular science, and 
made to serve as a beautiful proof of animal intelli- 
gence. This kind of arbitrary misrepresentation of 
the plainest facts can certainly lay no claim to any 
scientific value. 

How, then, do ants build their nests? They adapt 
themselves to given situations and prudently take 
into account the various circumstances. When in 
spring a gentle, warm rain begins to render the dry 
soil soft and manageable, immediately the earth- 
working ant species are kindled with new zeal for 
building. These assiduous little animals will then 
sally forth from their nests by hundreds and place 
pellets of earth upon pellets to build new galleries and 
chambers, availing themselves of blades of grass, 
twigs of heather, pieces of leaves and other natural 
props as pillars or vaults. For the same purpose such 
auxiliary materials are also first dragged to the spot. 
In this connection the sanguine slavemakers give evi- 
dence of eminent skill in combining timber-work with 
masonry. 

It is especially remarkable with earth-working ants, 
that they accommodate their instincts to the changes 
of temperature and moisture. This could be ascer- 
tained constantly in glass nests, in which I kept under 
observation small colonies of Lasius niger, Tetramo- 
rium caespitum and Myrmica scabrinodis. If the mois 
ture of the nest became too great, the ants would set 
to work and pile up the earth in the shape of a dome 
perforated by innumerable openings, making it appear 
like a sponge; thus the evaporation of the water was 
facilitated. But when the moisture decreased too 


Architecture in the Animal Kingdom. 117 


much and the nest threatened to dry up, a change to 
the opposite was effected; the nest was constructed as 
flat and as low as possible with very few openings on 
the surface. The propriety of such proceedings is cer- 
tainly striking. Often enough the same can be 
observed also in nature, both in those species that 
build only domes of earth, and in those that build 
regular ant-hills. It is a fact even noticed by farmers 
and ascertained by myself repeatedly, that in dry and 
hot summers the hills of wood ants are lower and 
flatter than in moist and cold summers. The first way 
of building is for the purpose of reducing evaporation 
to the lowest limit and to offer to the hot rays of the 
sun but a small surface to shine upon; on the other 
hand, the higher and the more vaulted the hills are, 
the easier is the drainage in case of rains, and the 
greater are the evaporating. and heating surfaces. 
Indeed, ants would have to be very intelligent, if their 
own reflection should lead them to modify their nests 
so prudently. Yet, considering that the dome shape 
for the nest includes in principle these quasi-intelli- 
gent adjustments to the variations of temperature and 
moisture, which can be traced, in consequence, to the 
specific manner of building proper to the ants in 
question, it is clear that instinct and not intelligence 
is the guiding principle; for, the specific plan of the 
building is, no doubt, instinctive, as is acknowledged 
at least by scientific authors. In adjusting, however, 
their instincts to changing circumstances, the animals 
are influenced by sensitive perceptions and experi- 
ences, which are nothing else than the natural exercise 
of the same instinctive power of cognition, on which is 


118 Chapter III. 


based the specific mode of building in any species of 
ants. Why, therefore, introduce a foreign element, 
called intelligence, between this hereditary disposition 
and its changeable application? I should think it is 
far more simple and natural to account for the whole 
activity of an animal by one and the same principle. 
Unless you mistake instinct for mechanical automa- 
tism, it is by no means necessary to assume animal 
intelligence in order to explain the above phenomena. 

Let us now compare the architecture of ants with 
that of birds. There are several important differences. 
The nests of birds are more artistic! and regular; 
yet they are stamped with the unmistakable marks of 
monotony and uniformity within the same species, 
they are products of instinct in the strictest sense of 
the term. Moreover, as Altum? has admirably proved, 
the architecture of birds is a function of their 
breeding instinct. It begins at a certain stage 
of the development of this instinct; both reach 
their climax at the same time, and then grad- 
ually vanish together. On this account the nests 
for the first hatching in spring are, as a rule, built 
better than those later in the season. In this case 
practice does not form the master but the bungler.* 
With ants, however, architectural skill is found in the 
-workers throughout their life, it is suitably carried into 


1) By the way, this art has often been exaggerated. See ‘Die 
Baukunst der Voegel auf ihren wahren Wert zurueckgefuehrt,” in 
“Jahrbuch der Naturwissensch.”’ I (1885-1886), 198. 

2) “Der Vogel und sein Leben” (6th ed.), p. 163 ff. 

8) Something similar obtains also among the Coleoptera in the 
ingenious nest-building of the leaf-rolling beetle (Rhynchites betulae). 
See Wasmann, “Der Trichterwickler, p. 78 ff. 


Architecture in the Animal Kingdom. 119 


operation under the most different circumstances, and 
is even to a certain degree improved by sensitive 
experience, i. e., it is modified according to new per- 
ceptions. True, many birds, too, are able to adapt to 
changing circumstances’ the places and the materials 
they choose for their nests; nor can their instinct be 
called mechanical automatism, because its activities 
are governed by the sensitive cognition of the animal. 
Many species of birds change the place and the 
material of their nests according to locality; besides, 
the single individuals are not rigidly bound to a cer- 
tain material for building their nests, but they fre- 
quently employ scraps of paper, horse hair, cotton and 
other materials, which happen to come in their way. 
Those birds that build nests more or less open to view, 
instinctively avoid materials the coloring of which 
would set their nest into striking contrast with the 
surroundings. In this they are evidently governed by 
their power of sense-perception. Sometimes this pro- 
tective resemblance of the nest with its background is 
effected quite of itself, when birds use the material 
for their nests, which is usual and natural to them; 
but sometimes also unusual materials found by chance 
serve the same purpose. Thus I was informed by a 
friend:? “In Blyenbeck (in the northern part of Dutch 
Limburg) I had occasion to observe how chaffinches in 
building their nest ‘ingeniously’ imitated the greyish- 
white color of lichen which covered the tree on which 
they built. They used small scraps of paper, and thus 


1) Several examples of the same are found in ‘“‘Westfalens Thier- 
leben,” Vol. II. Besides also in Darwin’s posthumous essay on instinct 
(Romanes, “Evolution in the Animal Kingdom,” appendix). 

2) Rev. L. Dressel, S. J. 


120 Chapter III. 


they disguised their nest.’”* The visual resemblance 
between the white tree-lichens and the paper-scraps, 
which impressed the sensitive power of perception of 
those chaffinches explains quite naturally their seem- 
ingly intelligent proceeding. 

He, who concurs with Darwin? in attributing intel- 
ligence to birds when their actions are influenced by 
sensitive cognition, must credit ants with a still higher 
degree of intelligence; for it cannot be denied that 
birds, in building their nests, show far more specific 
uniformity than individual variability, whereas in ants, 
as a rule, the contrary is the case; with them the 
“psychic plasticity’ of the nest-building instinct is no 
doubt much greater. 

Yet the chief point of excellence, which distin- 
guishes the architecture of ants from that of birds is 
the number of uses to which it may be put. Birds 
build nests to serve as places for hatching their young 
only ; except during the pairing season, birds do not 
know their nests, nor does it ever occur to them to 
use them as dwelling-places. With ants, however, 
nests serve as permanent abodes for the whole family 
and often also for strangers of different species, which 
are hospitably received as guests. Finally, ants use 
their architectural skill for many other purposes. But 
before entering into particulars on this head, we would 
like to draw a comparison between the architecture of 
mammals and that of ants. 

In as far as the buildings of many mammals are 

1) Ch. Darwin, too, mentions already a nest of a chaffinch described 
by Hewitson, in which, instead of lichens, shreds of paper likewise 


had been employed (l. c., p. 417). 
2) L.c., p. 414. 


Architecture in the Animal Kingdom. 121 


not only nests but also permanent dwellings both for 
the parents and the young, they bear a closer resemb- 
lance to the nests of ants than to those of birds. 
Instances are plentiful; for the burrows of badgers, 
foxes and wild rabbits are well known. In some cases 
the same buildings serve also for storing provisions, 
as is the case with German marmots and moles. These 
buildings, therefore, by their greater universality of 
purpose, are more similar to ant-nests than to bird- 
nests. Another point of similarity with the former 
is in this, that they show, with some species at least, 
a greater individual variability and less specific uni- 
formity than the latter. Nevertheless, in all these 
respects ant-nests by far excel the buildings of mam- 
mals. The very highest vertebrates, the anthropoid 
apes, scarcely manifest a trace of building instinct or 
of -its intelligent application, unless you wish to 
mention the sleeping-places somewhat resembling 
regular nests, which Orang-Utans* are wont to build 
on trees. Although the brain of apes most resembles 
that of man, yet the most “intelligent” architects 
among mammals are found, not among the apes but 
among the rodents, which in development of brain 
are far inferior. Beavers are the.only higher animals 
whose architecture can bear comparison with that of 
ants. 

The buildings of beaver-families consist of an 
underground chamber and burrow resembling those 
of other mammals, and of a so-called “lodge.” The 
manner in which the latter is built was ably described 

1) See Buettikofer, ““Zoologische Skizzen aus der Niederlaendischen 


‘Expedition nach Central-Borneo” (Compte rendu du 3me Congrés 
international de Zool.), p. 224. 


122 Chapter III. 


by Friedrich in a recent publication.t The beaver- 
lodge is nothing but an accumulation of brushwood 
above the opening of the underground chamber, which 
is the real center of the whole dwelling. Wherever 
beavers are living in colonies and when circumstances 
favor the full development of their instinctive skill in 
building, they construct their well-known dikes? to 
dam the water, and sometimes they even build canals 
for the transportation of timber. Although these 
works are the result of the co-operation of several 
families, yet each pair works only for its own pur- 
poses; there is never any division of labor like that 
in ant colonies. Of course, observers like Lewis H. 
Morgan,* who mistake for intelligence every action 
due to sensitive cognition, discover many proofs of 
high intelligence in the doings of the American 
beavers. However, this so-called “free intelligence” is” 
nothing else than the power of adapting their buildings” 
to the changes of situation. This power is possess 
also by ants in at least an equal degree. If Mo 
and Romanes,‘ e. g., regard it as an infallible proof” 
of the intelligence of beavers that they regulate the 

level of their ponds by widening or narrowing “the 

orifices of their dams as the case may be,” they should 

admit the same in ants, which regulate the degrees of 

moisture and temperature of their nests by changing 










1) “Die Biber an der mittlern Elbe” (Dessau, 1894), p. 20 ff. 

2) According to Friedrich they also occur at some places on the 
banks of the middle Elbe, although indeed on a smaller scale; there- 
fore they are due to an instinct common both to the European and 
the American beaver. 

8) “The American Beaver and his Works” (Lippincott & Co., 
1868). 

4) “Animal Intelligence” (6th ed), p. 377 ff. 


Architecture in the Animal Kingdom. 123 


their domes accordingly; and, just as beavers use 
their architectural skill not only in building nests but 
also in constructing dams and canals, so ants use theirs 
for far more various purposes. Those who concur 
with Romanes in maintaining that “the adaptations 
of pure instinct have reference only to conditions that 
are unchanging’, and that meeting of continual vari- 
ation of conditions cannot be accounted for but by 
intelligence, must, indeed, attribute to beavers a con- 
siderable degree of individual intelligence, but no less 
to ants. However, this conception of instinct and 
intelligence is untenable. Even Romanes is loath to 
ascribe to beavers such high psychic faculties, yet 
with his false notion of intelligence this conclusion is 
unavoidable. If beavers in modifying their buildings 
are guided by their own reasoning, they must be like- 
ise credited with an intelligent knowledge of the 
inciples of their architecture; for the former without 
atter is impossible; thus, we have, instead of 
building instinct, the highest order of human, 
rchitectural intelligence! This is evidently untenable. 
Those, however, who explain the architectural instinct 
of beavers from the hereditary disposition of their 
sensitive cognition and appetite, are able to explain 
from the same principle any given modification of their 
architecture, without resorting to “animal intelli- 
gence.” 


5. Other Purposes for Which Ants Employ Their 
Architecture. 








Some, ants having populous colonies often establish - 
temporary stations at the foot of trees and bushes, 


it ee Ae i sy 


124 Chapter ITI. 


where they visit their plantlice and scale-insects to 
“milk” them by caressing them with their feelers. A 
few European ants, namely Formica rufa, pratensis 
and Lasius fuliginosus build regular streets, clearing 
away from their path all vegetable growth to a dis- 
tance of sometimes from 20 to 50 m.t_ These streets 
lead from their nests into woods and bushes and thence 
branch off to the pasture-grounds of their “cattle.” 
Other ants, in particular Lasius niger and Cremas- 
togaster scutellaris build covered roads or tunnels of 
earth, by which their nests communicate with trees 
and bushes that are inhabited by aphides or scale- 
insects. These they occasionally surround with earth- 
ramparts, in order to keep them together and to pro- 
tect them from other ants by whom they might be 
coveted. Such a “plantlice-pavilion,’ an earth con- 
struction the size of a hazel-nut, is in my collection; 
it was built by Myrmica scabrinodis at the top of an 
oak-twig in the neighborhood of Exaten. Other ants, 
e. g., the African Dorylus species, dig subterranean 
tunnels, where they go for their prey, consisting 
chiefly of insects and worms. The harvesting ants of 
Southern Europe, Western Asia, Northern Africa, 
America and India establish granaries in their nests, 
where they store up their provisions for winter or sum- 
mer. The Afta of tropical America, feeding on mush- 
rooms, use a number of subterranean chambers as vege- 
table gardens and hot-houses, in which the mushrooms 


1) Among foreign ants there ate especially the larger species of 
the American leaf-cutting ants (Atta) which, according to Belt, Brent 
and Forel, build similar roads, but frequently of a still more consider- 
able length and breadth. The same ant-roads we find in North America 
with ants of the group of F. rufa, especially with Formica exsectoides. 


Architecture in the Animal Kingdom. 125 


are grown.’ Ants employ their building skill also to 
protect themselves against enemies. They raise ram- 
parts and barricades to keep off foreign invaders; and 
unwelcome visitors which cannot be got rid of in any 
other way, are simply walled up with earth, and are 
thus kept at a distance. Thus, in one of my observa- 
tion nests of F. sanguinea a salamander introduced 
by me was in a short time entirely walled in. The 
slaves (F. fusca), past masters in the art of building, 
were most zealous in this work. An occurrence far 
more amusing took place in a nest of Lasius flavus, 
to whom I had given a Lomechusa strumosa as guest. 
The small, yellow ants were not at all pleased with 
the unwieldy fellow, and tried to get rid of his impor- 
tunity in the following droll manner. From all quar- 
ters they brought together pellets of earth and heaped 
them up on the back of the unfortunate beetle, until 
nothing was to be seen of him but the tips of his 
feelers.” 

Ants are even supposed to have their cemeteries 
and burial-places. Superficial observers have circu- 
lated many fables on this subject. In the book of a 
certain Reverend White (Ants and their Ways, Lon- 
don, 1883), I found a touching story by Mrs. Lewis- 
Hutton, of Sidney, which is really too characteristic of 
this kind of natural history to be passed over in silence. 
One of her children had sat down on an ant-nest and 
had been assailed by the enraged inhabitants. At the 


1) Moeller, “Die Pilzgaerten einiger suedamerikanischer Ameisen’”’ 
(Jena, 1898), and Forel, “Zur Fauna und Lebensweise der Ameisen 
im Columbischen Urwald” (Mitteil. der Schweiz. Entom. Gesellsch.,”’ 
IX, 9th issue), p. 406. 

2) “Die internationalen Beziehungen von Lomechusa strumosa,”’ 
in “Biol, Centralbl.,” 1892, p. 653. 


126 Chapter III. 


cries of her child the mother ran to the spot and killed 
a few score of ants. A short time after she saw the 
corpses surrounded by a number of their companions. 
The burial ceremonies began. A deputation of ants 
was despatched to the nest to fetch the train of 
mourners. They marched in due order two by two to 
the scene of disaster. They took up the corpses, 
marched slowly in procession to a sandy place in the 
neighborhood and buried them one by one. A few of 
the gravediggers which tried to escape this doleful - 
duty by flight, were pursued by the other ants, over- 
taken and summarily sentenced to death. The sen- 
tence was immediately carried out, and the criminals 
were all interred in a common pit. The said lady 
maintains to have witnessed similar proceedings more 
than once. Gerstaecker in his “Report on the scientific 
results obtained by Entomology during the year 1861” 
mentions this burial story (p. 156) with the follow- 
ing ironical remark: “To render the mystification 
complett, nothing more was lacking than a funeral 
sermon held by one of the ants.” Strange to say, 
Perty? attempts to defend the imaginative lady against 
Gerstaecker by saying: “There seems to be some 
truth in it, anyway, for Dupont also maintains that 
ants have common graveyards at some distance from 
their buildings, whither they carry their dead.” 
Ernest André? was far more correct about those burial 
ceremonies of ants in calling them phantastic misrep- 
resentations of the commonest occurrences. It seems 
scarcely possible, that such an anecdote should see 


1) “Seelenleben der Thiere” (2d ed.), p. 328, 
2) “Les fourmis” (Paris, 1885), p. 176. 


Architecture in the Animal Kingdom. 127 


the light in a highly scientific publication, in the Trans- 
actions of the Linnean Society of London (1861). 
We are scarcely able to understand, why George 
Romanes, who incorporated the Australian burial 
anecdote in his work “Animal Intelligence,” did not 
even in the sixth edition (1895) entertain any doubts 
about it, but continued to regard it as_ sufficiently 
authentic. Something, to be sure, was true in the 
whole affair; but this “something” is confined to the 
fact that ants remove their dead from their nests to a 
certain place where they collect all their rubbish, and 
sometimes cover it with earth. These places, how- 
ever, would no doubt be far more correctly called 
“dumping grounds” than graveyards; for they are 
nothing but places where they deposit everything 
that displeases their instinct of cleanliness.1 These 
phenomena have nothing to do with “Chinese venera- 
tion for the dead,’* as can be easily ascertained in 
artificial observation nests. In these nests the dryest 
spots farthest away from the interior of the main nest 
are used for collecting the refuse. In my large obser- 
vation nest of F. sanguinea the place bearing the namg 
“refuse nest” is the one in which the corpses of dead 
ants, remnants of dead flies, wings of dismembered 
dragon-flies, empty cocoons of ants and other rubbish 
is finally stored, just because the ants wish to get rid 

1) Forel, “Ameise und Mensch oder Automatismus und Vernunft.” 

2) W. Marshall, in his ‘“‘Leben und Treiben der Ameisen” (p. 26), 
says, ‘the American forms seemed (as regards the treatment of their 
dead) to be more affectionate than those of the old world.” We should like 
to know why? At least nothing of the kind follows from the observa- 
tions, which he mentions, of McCook. See “Die Honigameise des 
Goettergartens” (“Stimmen aus Maria-Laach,” XXVII, 1884, 282), 


where we have dwelt more at length on the “affection” of the Ameri- 
can honey-ants. 


128 Chapter III.. 


of such disagreeable objects. It is utterly useless to 
embellish the life of ants with fabulous anecdotes like 
that Australian burial story. The cold facts are inter- 
esting and wonderful enough. 


6. Is the Architecture of Ants Guided by Intelligence? 


The building instinct of ants proves to be such a 
universal faculty, and its application to various pur- 
poses is in many cases seemingly so intelligent, that 
we are finally confronted by the question: Why 
should we not call this an intellectual faculty? The 
following discussion will probably throw some light 
on the subject. 

Would it not be a proof of intelligence, if ants, 
not themselves provided with spinning glands, 
employed their larvae for manufacturing threads, by 
means of which they build a nest of leaves? Accord- 
ing to W. D. Holland’s observations this is done by a 
large, reddish-yellow ant of Eastern Asia called 
Oecophylla smaragdina, whose nests he studied in 
Ceylon.t. With their mandibles the ants first bring into 
the proper position the leaves to be connected and keep 
them there; then others approach in large numbers, 
each carrying a larva in its mouth, with which they 
begin to move across the leaves from side to side. 
Wherever the mouth of the larva touches the leaf, a 
thread appears sticking to the leaf. This process is 
continued, until the leaves are attached to one another 
at their edges by a firm tissue of threads, and finally 
a viscous, paper-like stuff is formed consisting of 
innumerable threads crossing each other in all direc- 


1) E. E. Green, “On the habits of Oecophylla smaragdina F.”’ 
(“Proceedings of the Entomological Society of London,”’ 1896, p. IX.). 


Architecture in the Animal Kingdom. 129 


tions. These ants use their larvae as “spinning 
wheels,” not only for building their paper-nests, but 
also, according to Holland, for protecting their nests 
against the invasions of small ants, with whom they 
are in continual warfare. Around the trunk of the 
tree containing their nest they sometimes put a belt 
a foot broad, formed of threads, which serves to 
entangle the small ants and prevent them from climb- 
ing the tree. In manufacturing this protective tissue 
the ants come forth from their nest, each carrying a 
white pellet in its mouth, and move to and fro on the 
trunk. Upon closer inspection it was found that those 
little white lumps were again their larvae! 

These curious phenomena needed to be corrobo- 
rated by further investigation. Forel in his “Die 
Nester der Ameisen” (p. 20) already called attention 
to the fact that the mandibular glands of Oecophylla 
-- are strongly developed and possess large and numer- 
ous cells. From the analogy with other ant species 
(Cremastogaster, Dolichoderus, etc.), which secrete 
a certain glue from their mandibular glands for manu- 
facturing their paper-nests, it might seem probable, 
that the spinning material of Oecophylla proceeds 
from the ants’ mouth. But, on the other hand, Pro- 
fessor Chun has recently shown in his splendid book 
“From the Depths of the Ocean,’’! that the spinning 
glands of the Oecophylla-larvae are far more developed 
than those found in other larvae of ants. We must. 
conclude, therefore, that Mr. Holland’s statements 
were quite exact, and that the spinning glands of the 
larvae, not the salivary glands of the ants themselves, 


1) “Aus den Tiefen des Weltmeeres,”’ 2d ed., Jena, 1903, p. 129. 
9 


180 Chapter III. 


deliver the threads employed in the marvelous archi- 
tecture of Oecophylla. 

In adopting this supposition, that ants employ 
their own children as a kind of “spinning wheel,’ we 
are confronted by the extraordinary fact that animals 
make use of an instrument, other than any bodily 
organ, for building and defending their nests, an 
occurrence unparalleled in the whole animal kingdom, 
even among higher animals. But can we account 
for the proceedings of this Indian ant on the score 
of intelligence, that is to say, of her own, individual 
reflection? Just as little as in the case of the other 
specific arts and talents of animals; for they are all 
the hereditary property of certain species, not invented 
or learnt by independent individuals. And, therefore, 
also the spinning talent of Oecophylla, even if it is 
done by means of the larvae, is due to hereditary 
instinct, not to the individual intelligence of the animal. 

To obtain proofs for attributing the architecture 
of ants to their own intelligence we sliould have to 
look for instances, in which, in consequence of indi- 
vidual experience and reflection, these animals modify 
their innate instincts in such a manner as to invent 
new means of accomplishing their purpose. This is 
the third form of independent learning, which, as we 
have shown in a former publication, is a real proof 
of the intelligence of the learner. We must, there- 
fore, examine, whether the building activity of ants 


1) In the chapter on the different forms of learning in ‘Instinct 
and Intelligence” (Herder, St. Louis, 1903). A more detailed discussion 
of this point will also be found in our publication, ‘Die psychischen 
Faehigkeiten der Ameisen” (Zoologica,” 26th issue, Stuttgart, 1899), 
pp. 82-114. 


Architecture in the Animal Kingdom. 131 


contains any facts belonging to this class of psycho- 
logical actions.- 

Patrons of animal intelligence have repeatedly 
maintained, that ants build bridges with the intelligent 
purpose of overcoming obstacles placed in their way. 
Even a century and a half’ ago it was noticed by 
Cardinal Fleury, that when he tried to keep ants from 
climbing a tree by smearing it with bird-lime, they 
gradually covered the bird-lime with earth and thus. 
paved a road across. He informed the famous Réau- 
mur of this observation, who immortalized it in his 
Mémoires pour servir a l’histoire naturelle des Insectes 
(1734-1742). And because ants have not become 
weaker-minded since the days of Réaumur, many 
other friends of nature had the occasion, from that 
time on, to observe and.to record similar “bridges” 
built by these insects. One of these reports, best 
known in recent times, is the following.t Professor 
Leuckart in Giessen (Germany) wishing to keep ants 
from frequenting a certain tree, surrounded it with a 
cloth soaked in tobacco-juice. The ants above the 
cloth upon meeting the impediment turned back and 
after awhile let themselves drop to the ground from 
the branches, but those which were ascending to visit 
the aphides, after-having in vain tried to cross the ill- 
smelling cloth, descended and after a short time were 
seen coming back, each with a pellet of earth in its 
mouth. These they pasted over the tobacco-juice, and 
soon a passable road was constructed. William Mar- 
shall recording this observation of_Leuckart in his 
“Leben und Treiben der Ameisen” (p. 40) adds the 


1) Buechner, “Geistesleben der Thiere,” p. 116, 


182 Chapter III. 


following momentous reflections: “All the philos- 
ophers together of ancient and modern times, and all 
the theologians moreover, will not impose upon me 
by asserting that we have to do here with the action 
of an unreasonable creature. If this is instinct, then 
the invention of the steam-engine is instinct, too! No, 
both mean a clever profiting by given circumstances, 
due to reflection !” 

Many an unwary reader may, possibly, be over- 
whelmed by this spirited appeal to the steam-engine 
on the part of Marshall. However, if we do not allow 
ourselves to be imposed upon by the boldness of his 
oratorical flight, we shall arrive at different results, 
without being exactly philosophers or theologians. 
We can observe any day, that on the part of the ants 
ill-smelling or sticky objects are simply covered with 
earth, if they cannot be removed from the nest. 
Nobody will be inclined to maintain seriously, that 
ants, by so doing, make use of any “intelligent gift of 
invention” transcending their power of instinct. Out- 
side of their nests also, and governed by the same 
instinct, they occasionally adopt the same procedure. 
Now, in the above mentioned case the ants found, that 
the road which led them to their aphides on the tree, 
had been covered with an ill-smelling, sticky substance. 
What was more natural than to fetch pellets of earth 
and to clear a passage by a method so familiar to their 
instinct? Hence, we are justified in drawing the fol- 
lowing conclusion: That by this pretended “bridge- 
building” these ants have given infallible evidence of 
reflection, inventive genius, and intelligence, is a state- 
ment worthy only of popular, uncritical psychology. 


Architecture in the Animal Kingdom. 133 


One thing, however, is made evident by these 
and similar observations, viz.: that ants are not mere 
reflex machines, but beings endowed with sensitive 
cognition and appetite, and with the power of employ- 
ing in the most various manner their innate, instinc- 
tive faculties and abilities under the influence of 
different sense-perceptions. And just on this account 
it is altogether superfluous to admit “animal intelli- 
gence”; for, the complex representations of sensitive 
cognition, as we have shown in the above example, 
afford a simpler and better explanation of whatever 
is not mere fiction in those supposedly intelligent 
actions of animals.? 

Another example of bridge-building, which, by the 
way, is merely vouched for by a Mr. Theuerkauf in 
Buechner’s “Geistesleben der Thiere” (p. 117), is 
still less corroborative of ant intelligence than the 
former. In this case the ants used a different means 
for bridging over a circle of tar smeared around a 
tree. The ants were descending from the top; on — 
arriving at the obstacle some stuck fast, others 
returned to fetch plantlice from the twigs; they put 
them oh the tar and thus constructed a bridge. Sir 
John Lubbock? remarks in explanation of this story, 
that he had his doubts as to the interpretation of the 
fact. “Is it not possible that, as the ants descended 
the tree, carrying the aphides, the latter naturally 
stuck to the tar, and were therefore left there? In 
the same way I have seen hundreds of bits of earth 


1) Cf. on this point, “Instinct and Intelligence in the Animal King- 
dom” (1903), p. 109 and 137 ff., where we have shown, that not even 
higher animals may be credited with formal consciousness of purpose. 

2) “Ants, bees and wasps’? (London). 


134 Chapter III. 


deposited on the honey, with which I fed my ants.” 
In fact, only trivial observers could maintain that in 
this case the ants had intentionally employed their 
aphides as bridge-building materials. The correct 
explanation might rather be the following: the ants, 
becoming uneasy about their precious aphides living 
on the tree, tried to save them by carrying them down. 
By this attempt, however, the aphides literally “got 
stuck in the mud.” It was, no doubt, merely by 
chance, that the aphides adhering to the tar formed 
a sort of bridge for the ants. 

These two famous stories are, therefore, far from 
furnishing any evidence in favor of ant-intelligence. 
We have to investigate other examples to determine 
whether or not ants are able by “reasonable reflection” 
to invent new means for fulfilling their designs. 

Sir John Lubbock? has made a number of experi- 
ments with ants in order to test their intelligence ; some 
of the more important only can be mentioned here. 
For the ants of a nest of Lasius niger he arranged a 
bridge made of a piece of straw or a slip of paper, by 
which they could get at their larvae. After the ants 
had become sufficiently familiar with this pathway, 
he slightly moved the bridge, ‘“‘so as to leave a chasm, 
just so wide that the ants could not reach across. They 
came and tried hard to do so; but it did not occur to 
them to push the paper bridge, though the distance 
was only about one-third inch, and they might easily 
have done so.” Another experiment he relates as 
follows: “I suspended some honey over a nest of 
L. flavus at a height of about half an inch, and accessi- 


1) “Ants, bees and wasps,” Chap. IX. 


Architecture in the Animal Kingdom. 136 


ble only by a paper bridge more than ten feet long. 
Under the glass I then placed a small heap of earth. 
The ants soon swarmed over the earth on to the glass 
and began feeding on the honey. I then removed a 
little of the earth, so that there was an interval of 
about one-third of an inch between the glass and the 
earth; but, though the distance was so small, they 
would not jump down, but preferred to go round by 
the long bridge. They tried in vain to stretch up from 
the earth to the glass, which, however, was just out 
of their reach, though they could touch it with their 
antennae; but it did not occur to them to heap the 
earth up a little, though, if they had moved only half 
a dozen particles of earth, they would have secured 
for themselves direct access to the food.” It is evident 
from this, that the ants had not the least idea of em- 
ploying even this simple means. In all his experi- 
ments Lubbock obtained entirely negative results. 
There was no indication of the much-vaunted intelli- 
gence of ants. Strange to say, William Marshall has 
not mentioned these experiments of Lubbock in his 
“Leben und Treiben der Ameisen,” although they 
could not have been unknown to him, as he translated 
them himself into German; perhaps it was, because 
the results did not fit in with his enthusiastic praise 
of the “reflective faculty” and “inventive genius” of 
these animals. 

This latter experiment of Lubbock was repeated 
of late by Albrecht Bethet in a somewhat different 
form. Over a well-frequented pathway of Luasius 


1) “Duerfen wit den Ameisen und Bienen psychische Qualitaeten 
zuschreiben?” (Bonn, 1898), p. 66. 


136 Chapter III. 


niger he fastened a strip of tin with some honey. 
After the ants had for a long time been allowed to 
visit the honey, the strip was gradually raised by a 
screw, until from their pathway the ants could no 
longer get at the honey. Though it would have been 
easy enough to heap up a little earth under the strip of 
tin, it never occurred to the ants to do so; the honey 
remained beyond their reach. This experiment, there- 
fore, had the very same results as Lubbock’s, namely, 
that the ants were not capable of forming the simplest 
intelligent conclusion, which would have led them to 
employ their building skill for the purpose of getting 
at the honey. 

I may add here a few observations and experiments 
of my own. Since it might be objected against Lub- 
bock’s results, that he took for his experiments some 
ant-species “little endowed with intelligence,” namely, 
Lasius and Myrmica, I chose the most intelligent ants, 
namely, Formica sanguinea* and her allied slaves as 
subjects for experiments, of which only a brief extract 
is here presented. 

In the front-nest of my above mentioned observa- 
tion-nest (see p. 23) a piece of wood formed a com- 
modious bridge, over which the ants could pass to the 
rim of the glass and thence into the top-nest. By their 
earth-constructions in the front-nest the ants had 
gradually lowered the bridge, so that the distance 

1) Bethe infers from this experiment that ants do not even possess 
sensitive perception and cognition. This inference is too far-reaching, 
and is owing to his mistaking intelligence for sensitive cognition. See 
“Die psychischen Faehigkeiten der Ameisen,” p. 73. 


2) Forel also (‘‘Fourmis de la Suisse,” p. 443) states that F. 
sanguinea deserves the palm for intelligence, 
’ 


Architecture in the Animal Kingdom. 187 


between the top-end of the bridge and the cork of the 
glass was about 2 cm. When exposed to the sun the inside 
of the glass was generally covered with moisture, and 
the ants found great difficulty in passing the intermedi- 
ate space and in getting into the top-nest. Although 
this state of affairs lasted for weeks, and the ants con- 
tinued to encounter the same difficulties, it never 
occurred to them to connect the broad, upper end of 
the stick with the roof of the front-nest by a bridge 
of earth. They connected the sides of the stick with the 
glass by a wall of earth; gradually they also covered 
the whole glass-wall with pellets of earth to protect 
themselves against the rays of the light; but they 
never built a bridge at the critical spot, where one was 
evidently needed and of the greatest importance. The 
pellets of earth accidentally fastened there were, on 
the contrary, continually thrown down by the ants 
that crawled up the road. Thus exactly that spot 
over which the ants, if they were endowed with any 
power of thought and reflection, would be expected to 
build a bridge, was left slippery and smooth, and was 
still the same after half a year. 

In 1884 I repeatedly made the following experi- 
ment: In a tiny vessel I suspended some honey or 
ant-larvae over a nest of F. sanguinea, contained in a 
large “crystallisator”,’ so that the ants could touch the 
vessel with their antennae only, but could not reach 
it except by a very circuitous route. It should have 
occurred to them to heap up a little earth or some 
pieces of wood underneath the vessel, in order to form 
a “bridge” leading directly to the wished for goal. 


1) A low, round glass bowl, covered by a glass plate. 


138 Chapter 111, 


But neither the sanguineas nor their slaves (F. rufa 
and fusca), living in the same nest, ever hit upon this 
obvious method, although it would have sufficed to 
raise the surface of their nest at the spot in question 
just by 1 cm.! 

A more wonderful result was obtained in another 
experiment on the same nest of sanguinea. On June 
16, 1884, I filled a large watch-crystal with water and 
in the center upon a kind of island I placed a little 
shell filled with ant-cocoons previously taken from the 
same colony. This artificial pond with its island was 
then introduced into the nest. The ants soon noticed 
the cocoons and stretched out their feelers towards 
the island; but getting into the water at every attempt 
to approach, they retreated again and again. I 
began to think they would never be able to overcome 
the difficulty, when suddenly a sanguinea began to 
throw into the water pellets of earth, bits of wood, 
dead ants and similar solid materials. Others followed 
her example and they soon had built a road over the 
water! In the space of an*hour, counting from the 
minute I started the experiment, they had fetched all 
the cocoons from the island by means of this “floating 
bridge.” The very last cocoon having been secured 
by the ants, one of them returned to the island and, 
finding it empty, she squatted on her haunches, passed 
the spur of her fore-feet through her mouth and then 
combed her feelers with the spur, sitting there for 
several minutes in a most provoking attitude, as if she 
were saying to me: “Ah, my dear, who has won the 
game now?” Js this fact not a staggering proof, 


1) This very last instance, as many others in this translation, was 
added by the author from his original notes. 


Architecture in the Animal Kingdom. 139 


that at least the sanguineas are endowed with a good 
quantity of reflective power and of intelligent con- 
sciousness of purpose? 

In order to answer this question I tested the above 
observation by the following experiment: After some 
time I again placed the same watch-crystal filled with 
water in the nest, but this time without island and 
without cocoons. Now, supposing that the ants in 
the previous case had really intended to build a bridge 
for the sake of getting at the cocoons, there was no 
longer any reason for them to repeat the procedure. 
However, this second time also, after several had 
accidently got their feet wet, they soon started to fill 
up the lake. Although this time there were no cocoons 
to be obtained, nor any island in sight,’ yet they again 
covered the water with all kinds of materials in the 
same manner and almost in the same space of time as 
they had done before. Hence, we are allowed to con- 
clude, that even the first time the ants had not intended 
to build a floating bridge, but only to get rid of the 
disagreeable moisture that barred their way. If, there- 
fore, we maintained, that in the first experiment the 
ants had by intelligent fore-thought invented a means 
for regaining possession of their cocoons, we would 
be guilty of uncritically humanizing the brute. 

From all the observations made and noted down for 


1) I wish to lay emphasis on this circumstance, because Prof. 
Charles Sajé (in “Prometheus,” 1899, No. 486, p. 284) believes the 
ants had hoped to find some treasure on the island this time also. The 
ants could easily notice from the margin of the watch-crystal, that 
there was no island. The eyes of F. sanguinea being rather large and 
sharp and capable of distinguishing the form of small objects at a 
distance of several centimeters, the ants could undoubtedly see that 
there was no island, 


140 Chapter III. 


the last twenty years, I could record here still many 
an iftteresting occurrence, which, like the above exam- 
ple, impresses a superficial observer as an intelligent 
action. Yet, closer examination invariably proves 
that such facts are accounted for much more easily 
and naturally by the instinctive combinations of sense- 
representations ; therefore, no “ant-intelligence,’ and 
in fact no “animal-intelligence’ at all is required. 
Indeed, the higher mammals ranking next to man 
in brain development are far from supplying more con- 
vincing proofs of “intelligence” than ants. In them 
also the whole process of cognition is confined to the 
mere connecting of sense representations and sense ex- 
periences according to the inborn laws of instinctive 
association of representations, which ordinarily regu- 
late their lives. The psychic endowments of dogs 
and monkeys go no farther. Unless a dog has been 
specially trained, it never occurs to him to open a 
door, the knob of which he is unable to reach, by 
fetching for instance a foot-stool to gain a higher 
level; he may have seen children, his play-fellows, 
doing the same thing a hundred times; the relation 
between means and end, though so natural and obvious 
in this case, will forever remain hidden to the canine 
soul. Hence the dog is not a whit more intelligent 
than the ants, that failed to notice, that a little heap 
of earth would have sufficed to secure them an easy 
passage to the honey suspended in a saucer above their 
nest. 
Neither do apes possess the power to invent by 
their own reflection new means of accomplishing their 
end. Even these highest mammals are confined ex- 


Architecture in the Animal Kingdom. 141 


clusively to the instructive association of sense-repre- 
sentations. We have shown this in our first chapter, 
where we discussed the wars that take place in the 
animal kingdom, and pointed to the fact that apes are 
unable to invent the simplest weapons and implements 
even. The same holds good as to the use of fire. If - 
a troop of apes in the forest hits upon the remnants 
of a fire lighted by the hands of man, they will cer- 
tainly gather around it and enjoy the comfortable 
warmth. But it has never, hitherto, occurred to any 
ape to supply it with fuel.t And yet it would be such 
a simple and natural combination of representations, 
requiring but a low degree of intelligence. Why do 
apes, in spite of the “high plasticity” of their quasi- 
human brain, never hit upon such a simple means? 
Because they possess no spiritual soul and therefore no 
intelligence. ‘The “plastic neurozymic activities” of 
the simian brain are essentially different from human 
intelligence; like those of ants and all lower animals 
they prove to be functions of mere sensitive instinct. 
It is wrong, therefore, to describe ants as instinctive 
automatons, in order to safeguard the intelligence of 
the higher animals. The psychic actions of all animals 
are due to automatism, as far as they are unable to 
attain the level of reasonable reflection and free self- 
determination. For the rest, however, there is no 
question of automatism either with lower or with 
higher animals, because it is sensitive cognition and not 
mere reflex activity, which prompts them to act. It 
is true, that instinctive actions have a certain auto- 

1) Cf. Tylor (in Ranke, “Der Mensch,” II, 1st ed., 486) and 


Charles E. v. Baer (in Stoelzle, “Karl E, v. Baer und seine Weltan-. 
schauung,” pp. 304, 314). 


—:'142 Chapter ITT. 


matic character, inasmuch as they are, to a certain 
degree, predetermined by the natural constitution of 
the animal. However, inasmuch as they are governed 
by sensitive cognition and appetite and, therefore, 
capable of more or less modification within the prede- 
termined natural limits, they are not of an automatic, 
but of a spontaneous character. 

In conclusion, let us sum up the results of our 
comparative discussion on architecture in the animal 
kingdom. They are: Ants surpass all animals, both 
lower and higher, by the quasi-intelligent variability, 
the spontaneous self-determination and the power of 
suitable adaptation, manifested in their architectural 
skill. Nevertheless it is as certain of them as of any 
other animal, that they are not endowed with intelli- 
gence properly so called. 

_ This corroborates the views advanced in our dis- 
cussion on the different forms of learning. Ants are 
able to accommodate their buildings to the most varied 
conditions; hence they are able to “learn” how to 
modify their buildings according to given circum- 
stances. But this learning takes place only in so far as 
sense-experience gives rise to new combinations of 
representations; as soon, however,-as the modifica- 
tion of their activity would require intelligent reflec- 
tion, or the drawing of conclusions from former con- 
ditions to the present ones, then both ants and higher 
animals, without exception, are all at once incapable 
of further learning. This shows to evidence, that the 
doctrine of “animal intelligence” is utterly untenable. 
1) “Die psychischen Faehigkeiten der Ameisen,” pp. 82-114; ‘‘In- 


stinct and Intelligence in the Animal Kingdom” (Herder, St. Louis, 
Mo., 1903), Chap. VIII. 


CHAPTER IV. 
CARE OF THE YOUNG IN THE ANIMAL KINGDOM, 


1. A General View of the Breeding Instincts of 
Animals, 


UMAN community life naturally evolves from the 
family, which must always remain the founda- 
tion of the state. Animal societies have a similar 
origin and basis, though this similarity does not go 
beyond mere analogy. Wherever we meet permanent 
associations of animals, they are seen to depend, with 
very few exceptions, such as for instance the mixed 
colonies of ants, on the ties of common descent. The 
purpose of this social co-habitation is the preservation 
of the race and species. All other animal instincts are 
by natural law subordinated to that higher end, which 
is also the reason for the existence of social instincts 
of animals. 

One of the most important means for preserving 
the species is breeding, with the various instincts sub- 
servient to it. The different forms of breeding in the 
animal kingdom form one of the most interesting 
chapters of comparative animal psychology; in this 
place, however, we must confine ourselves to some of 
its more prominent features. 

With those lower animals, which reproduce with- 
out sexual generation by fission or by budding, there 
can be as little question of breeding instincts as with 

148 


144 Chapter IV. 


plants. The new being comes into existence already 
fully endowed with the power of subsistence according 
to the laws of merely vegetative nature, whether it be 
separated from the mother-organism, or remain united 
with it as a new part of a polyzoal colony. Here, 
therefore, it would be altogether useless to possess 
breeding instincts for the propagation of the species. 
Even among such animals as propagate through 
sexual generation, we meet with breeding instincts 
only where they are required for the preservation of 
the species; and the parents care for their off- - 
spring only in as far as it is necessary for that purpose. 
Within these limits, however, we find a wonderful 
adaptation of means to the end, and at times a mar- 
velous sagacity of animal instinct, which appears 
nowhere else to such advantage. 

But also nowhere else in the whole animal psy- 
chology are manifested so palpably the impotence of 
so-called animal intelligence and the unsoundness of 
the modern tendency of humanizing animal. life. How 
should the ephemera know by her “own intelligence,” 
that she may without any apprehension drop her eggs 
into the water? Does she perhaps still remember, that 
her “mother” once upon a time dropped her also into 
the water as an egg? Or has she perhaps by the 
study of zoology gained the knowledge that ephemeras 
need no hatching? According to Brehm’s psychology 
we ought to give the ephemera a thorough scolding 
for showing so little motherly love towards her dear 
offspring and for not caring for their welfare. But 
scientifically speaking, such a scolding is as nonsen- 
sical as if we were to blame an oak tree for bearing 


Care of the Young in the Animal Kingdom. 145 


acorns instead of pumpkins, or a hen for laying eggs 
instead of begetting live chicks. Ephemeras flit 
about over stagnant pools and drop their clusters of 
eggs into the water; pearl-flies carefully attach their 
eggs to a flimsy peduncle consisting of a sap hardened 
by exposure to the air; ichneumon-flies deposit their 
eggs in the body of a caterpillar by means of their 
ovipositor; gall-flies introduce theirs under the rib of 
an oak leaf, from which later on the gall-nut is to 
grow, serving both as dwelling and as provision store 
of the young larva; the blue-bottles place their eggs 
on putrefying flesh, whereas a certain species of wasps 
(Pompilus viaticus) glue theirs to the bodies of 
spiders which they paralyze by skilful thrusts of their 
sting without killing them, so as to enable the growing 
larvae to feed upon live flesh; the common cabbage- 
butterfly deposits her eggs on cabbages, the hawk- 
moth on poisonous spurges, the large clavicorn water- 
beetle (Hydrophilus piceus) weaves for its eggs an 
ingenious boat with a little streamer on top to float 
about on the surface of the water, whilst a smaller 
allied species (Spercheus emarginatus) carries its eggs, 
as many spiders do, in a bag attached to its abdomen ; 
the leaf-rolling beetle (Rhynchites betulae) cuts a 
birch-leaf in a manner implying a difficult problem in 
applied mathematics, and rolls it up into the shape of 
an ingenious funnel, in which it deposits its eggs; 
whilst Rhynchites pubescens saws a cradle for its eggs 
in the wood of an oak-twig, the ear-wig hatches its 
eggs like a hen, whilst Lomechusa strumosa, just like 
cuckoos, confides its brood to the care of ants; they 
all ae aes duty with equal prudence, but all too are 


146 Chapter IV. 


ignorant of the prudence of their actions, and uncon- 
scious of duty. Under the guidance of sensitive cog- 
nition and perception they follow the mysterious in- 
stinctive impulse arising from their organic develop- 
ment, which suggests to them the means necessary for 
preserving their species. But they do not understand 
the appropriateness of these means and need not reflect 
how to use them to advantage. 

Wherever in the animal kingdom the care of the 
young requires “family life,” i. e., a regular co-habita- 
tion of the parents and of the young, this task 
takes place only as far as the preservation of the 
species renders it necessary. The same organico- 
instinctive laws, to which breeding is subjected in 
general, also determine the existence and firmness of 
family ties as well as the extension of the family circle 
with different species of animals. There is no room 
for individual reason and liberty; and to postulate 
them is not only wholly superfluous, but also contra- 
dicted by innumerable facts. As birds associate in 
pairs only during the mating season for the preserva- 
tion of the species, so in building their nest and in 
hatching their young, the two mates co-operate like- 
wise only as far as is necessary for preserving their 
species; and the pairs remain together and in com- 
pany with their young no longer than the same purpose 
requires. Altum, in his excellent book “Der Vogel 
- und sein Leben”, has supplied us with a number of 
striking instances, proving how ridiculous and unten- 

1) We have so minutely proved this fact in the case of insects that 
live single, in our book, “Der Trichterwickler, eine naturwissenschaft- 


liche Studie ueber den Thierinstinct’” (Chap. IV. ff.), that there is no 
need of rereating the proof here. 


Care of the Young in the Animal Kingdom. 147 


able it is to apply to birds the notions of marital and 
maternal affection as it exists among men. As a 
matter of fact, there is no more “‘marital affection” in | 
the human sense of the term to be found with a loving 
couple of parrots than with spiders, where the smaller 
male must be on its guard not to be devoured by the 
larger female immediately after mating. And by 
devouring her “husband” the female spider sins as lit- 
tle against morals, as she acts conformably to them in 
carefully protecting and carrying along her egg-bag 
or in spinning a protecting web for her young. And 
the female cuckoo smuggling her eggs into nests of 
strangers acts as little against morals, as the foster- 
parents of the young cuckoos act conformably to 
morals in feeding and rearing these changelings. 
There is no room for reason and morality in the breed- 
ing instincts of animals; for they are exclusively deter- 
mined and regulated by the laws of organico-sensitive 
life. 

The same holds good for mammals, the anthropoid 
apes not excepted. As long as young dogs, cats, and 
apes need the care of their parents, they will not be 
forsaken. But no sooner are they old enough to shift 
for themselves, than their parents no longer know 
their once so “beloved” offspring. As the mates know 
each other only for sexual intercourse, so also they 
know their young only as helpless beings, whose 
behavior stimulates the nursing instinct of their parents 
. to action. As soon as this instinctive impulse ceases, 
then the mates and the young are completely estranged, 
having no regard for each other in the .relentless 
struggle for existence, for food and rut, just as if 


148 Chapter IV. 


they had never belonged together. This is a genenal 
law of nature, ascertained by science throughout the 
whole animal kingdom, setting at naught all the fine 
phrases and sentimental talk of Brehm on marital and 
parental love among animals. And this gush, the 
outcome of erroneous notions and misplaced sentiment 
is dubbed by thousands of its votaries modern animal 
psychology! 

Care of the young in its most primitive form is 
found among the Echinoderms, namely in a few spe- 
cies of star-fishes (Asterias Muelleri, rugispina, Cri- 
brella oculata).1 According to Perrier the female ani- 
mal, by bringing her arms near to the body, forms 
a kind of breeding cavity, in which the young, hud- 
dling together, are enclosed. In the different classes 
between the Echinoderms and the vertebrates care of 
the young assumes very different forms, which we are 
unable to discuss here. Of peculiar psychological 
interest, however, are those animals, among whom the 
males and not the females are entrusted with the care 
of building nests and rearing the young. The best- 
known example of this kind among fishes is the 
stickle-back (Gasterosteus aculeatus).* In this species 
the females are regular “cannibal stepmothers,” 
whereas the males are models of “affectionate fath- 
ers.” How ridiculous such facts are, when couched 


1) See H. Ludwig, “Sitzungsber. der Niederrh. Gesellsch. fuer 
Naturk.” (Bonn), 1896, 1st half, p. 104; besides in ‘‘Zoolog. Anzeiger,” 
1897, No. 534, p. 217 and No. 535, p. 237. 

*) Also among amphibias cases of male hatching have been ascer- 
tained. See Fr. Werner in “‘Verhandl. der Zoolog.-botan, Gesellsch. 
von Wien,” 1898, 1st issue, p. 11 ff. See also R. Wiedershein, 
“Brutpflege bei niederen Wirbelthieren” (Biolog. Centralbl. XX, 1900, 
Nos. 9 and 10). 


Care of the Young in the Animal Kingdom. 149 


in anthropomorphic language, goes without saying.’ 
Some instances of taking care of the young occur also 
among amphibias. The female of the Surinam toad 
(Pipa dorsigera) carries her young in the cavities of 
her dorsal skin; whereas in a frog species (Arthro- 
leptis seychellensis) inhabiting Central America the 
young hold on to the back of the male.? But quite 
universal and commonly known is the care bestowed 
by birds and mammals upon their young. Yet its 
highest perfection, connected with the most perfect 
form of community life in the animal kingdom, does 
not occur with the higher mammals, but with the 
social insects, in particular with ants. Here this degree 
of perfection is made possible by the organic division 
of the female sex into females proper and into nurses 
(workers) incapable of generation. And although 
these are not the mothers of the children they nurse, 
the psychic development of their breeding instinct 
reaches the greatest perfection in the whole animal 
kingdom. Before discussing, however, this aspect of 
- the breeding instinct of ants, we must first explain its 
connection with the laws of their organic develop- 
ment. 

The bodily differentiation of the members of an 
insect-state into classes and castes, their co-habitation 
in a common abode, their nest-construction, acquisition 


1) The following amusing quotation will do for the purpose: “The 
greatest danger threatens him (Mr. Stickleback) from the mothers of 
his own children. Eager to devour their own offspring, they ‘are con- 
tinually dashing in unison against the nest, in which the young are 
guarded by their watchful father, and but too often the latter pays the 
penalty of his polygamy” (Thilo, ‘‘Umbildungen an den Gliedmassen der 
Fische,” in ‘‘Biolog. Centralbl.,’”? 1897, 1st issue, p. 24). 

2) “Zoolog. Jahrb. Abth. fuer Systematik,” XII (1898), 89 ff. 


150 Chapter IV. 


of food, their whole life and activity have for their 
object the care of the offspring, and thereby the pres- 
ervation of the species. The animal colonies of bum- 
ble-bees, wasps' and hornets represent a lower stage 
of community-life, than the perennial colonies of 
honey-bees, ants and termites. In the case of the 
former the insect-families and the entire community- 
life have to be established anew every year by some 
hibernating female; but with the latter the original 
families last several years and often much longer, thus 
giving their community-life a character of stability 
and also of greater variety and perfection. 

The organic foundation of insect-states, as already 
shown above (p. 14), is polymorphism, or the separa- 
tion of the individuals into sexual animals and “neu- 
ters”? or workers. The prime duty of the former is 
generation, this being the direct means for preserving 
the species, whilst the latter perform all the work 
required for the welfare of the family, thus in their 
turn indirectly contributing to the same end. Without 
this appropriate division of labor insect-states would 
be impossible; and, as a rule, the more perfect this 
division, the more perfectly developed is the insect 
community.2 In bees, with whom the workers are 


1) According to H. v. Jhering (“Zoolog. Anz.,” Vol. XIX, 1896, 
No. 516, p. 449) a large number of the Brazilian social Vespidae 
(Polybia, Chartergus, etc.) form perennial colonies for several years, 
not annual ones as our native wasps. 

2) We have already in Chap. I, No. 2, referred to the fact, that in 
reality they are not, properly speaking, sexless. 

3) The greatest importance must be attached in this place to the 
differentiation between sexual individuals and workers. Thus e. g. in 
the annual colonies of bumble-bees there is a dimorphism of workers, 
and hence a more marked division of labor than in the perennial 
‘colonies of our honey-bee. (On the bumble-bees see esp. E. Hoffer’s 
excellent observations on the bumble-bee of Styria). Nevertheless the 
bee-states are more perfect than those of the bumble-bees on account of 
the greater difference between their workers and genuine females. 


Care of the Young in the Animal Kingdom. 151 


winged and not unlike the real females, polymorphism 
and, in consequence, community life is less differen- 
tiated than in ants, where the neuters are devoid of 
wings and,—with many species,—are again divided 
into different castes, namely, workers and _ soldiers. 
The greatest variety of bodily differentiation, how- 
ever, obtains in termites, which belong to the insects 
with imperfect metamorphosis. In ‘their case the 
larva resembles the imago and is transformed grad- 
ually with little change of outward appearance. There- 
by termites present the organic foundation for a still 
more abundant and variable division of castes, the 
formation of which may embrace not only sexual indi- — 
viduals on the one hand, and workers and soldiers on 
the other, but within these two categories again several 
different classes or forms.* 

Not even the most ardent defenders of modern 
animal intelligence would venture to attribute poly- 
morphism, which is the fundamental law in the con- 
stitutions of insect-states, to the “intelligence” of the 
animals themselves. It is evidently based on the 
hereditary laws of organic development. Just as it is 
not owing either to his own intelligence or that of his 


1) Cf. Hagen, ‘Monographie der Termiten” (“Linnaea Entom- 
ologica,” X-XIV); Grassi e Sandias, “‘Constituzione e sviluppo della 
Societa dei Termitidi,’”? Catania, 1898. (‘‘Atti dell’ Accademia Gioenia 
di Scienz. nat.,” 4, VI and VII); Wasmann, “‘Einige neue Termiten 
aus Ceylon und Madagaskar,” in ‘‘Wien. Entom. Zeitung,” 1893, 7th 
issue; “Neue Termiten und Termitophilen aus Indien” (‘‘Annali del 
Museo Civico di Stor. nat. di Genova,” 2, XVI, 1896, 613-630); 
“Termiten von Madagaskar und Ostafrika” (“‘Verhandl. der Senkenberg. 
Naturf. Gesellsch.,” XXI, 1897, 1st issue); G. D. Haviland, ‘“‘Observa- 
tions on Termites” (‘“‘Linnean Society’s Journal, Zoology,” Vol. XXVI, 
pp. 358-442). Dr. F. Silvestri “Ergebnisse biologischer Studien an 
suedamerikanischen Termiten” (‘‘Allg. Zeitschr. f. Entomol.,’” WIT, 
No. 9 ff.). 


152 Chapter IV. 


“mammy,” that a young rooster became a rooster 
and not a jackdaw, so no intelligence of the ant is 
responsible for the fact, that from the egg and the 
larva which she nurses, there emerges not a bee but an 
ant. All this is self-evident. Yet, the peculiar organic 
laws of development not only form the material 
of the breeding instincts, but also their directing 
principle. The breeding instincts of the different 
species are so well adapted to the hidden laws of 
organic growth, that no reflection or intelligence on 
the part of the animal, nay, not even the keenest human 
reason could ever succeed in inventing them. More- 
over, they are exercised by the workers completely and 
perfectly, without previous experience or instruction; 
they are innate in the animal, and grow with it, 
and when the young ant has reached the perfection of 
its organic development, they, likewise, are just as 
perfectly developed. Hence they must spring from 
_ the same source as the organic growth, that is to say, 
they spring from the organico-psychic laws of devel- 
opment of a given species, and have nothing to do 
with individual reason and free determination. As 
it is by organic development, that the male of an 
ant-species receives also the psychic endowment of a 
male, thus it is with the females and the workers. The 
distribution of psychic endowments in the different 
castes of an ant-state is regulated by the same laws 
as their bodily polymorphism. And this alone accounts 
for the fact, that within one and the same species the 
males are the most stupid members of the whole state, 
possessing the smallest brains, whilst the workers are 
endowed with many marvelous instinctive talents and 


Care of the Young in the Animal Kingdom. 153 


even surpass the females proper in perfection of 
instincts and brain development.t Those females, 
which are destined for generation, are provided by 
their organic development with perfect ovaries, whilst 
their brain and instincts are far less perfect. The 
workers, on the other hand, which on account of their 
small ovaries may be called undeveloped females, are 
compensated by a more perfect development of the 
brain and the instinctive endowments. Hence, the 
astonishing prudence displayed by the worker ants 
and their consequent social leadership are merely a 
function of their organic development. ‘This is the 
so-called “intelligence” and “intellectual life” of ants, 
viewed in the light of genuine science! 

We have thus far been considering the breeding 
instincts of ants from their organic side; let us, in° 
the subsequent discussion, turn our attention to their 
psychic aspect. 


2. Care of the Young among Ants. 


The hereditary disposition of the sensitive cogni- 
tion and appetite of animals, called instinct, has in the 
case of ants a wide range and great variety of actions, 
and especially so with regard to the breeding instincts, 
wherein ants surpass even the highest mammals. The 
instinctive disposition is no mechanical automatism, 


1) As to the peduncles of the ant brain, the significance of which 
with regard to psychic life we have,already pointed out in a former 
essay, Forel says: Les corps pédonculés sont énormes chez les ouvriéres 
du genre Formica, qui renferme les fourmis les plus intelligentes; et, 
chose trés remarquable, ils sont plus petits chez les femelles et 
beaucoup plus petits chez les males du méme genre (‘‘Fourmis de la 
Suisse,” p. 123). My own observations confirm Forel’s statements; see 
“Instinct and Intelligence in the Animal Kingdom,” p. 130 ff. 


154 Chapter IV. 


but is guided and influenced by various sensitive 
affections and perceptions. In bee-hives the eggs are 
simply deposited by the queens in the cells previously 
prepared by the workers, and the young bee-larva 
goes through the successive stages of development in 
one and the same cell. With ants breeding shows far 
greater variety and independence. The eggs laid by 
the queen are received by the workers and gathered in 
clusters of various dimensions. Then from all sides 
they are licked again and again with the utmost care, 
and begin to increase by the endosmosis of the nour- 
ishing juice.t This is the first stage in the rearing of 
the young in ant-communities. As soon as the egg 
has developed into a larva, there follows the second, 
the feeding and nursing of the larvae. When the time 
for entering the state of a pupa has arrived, the ant- 
larvae are carried by their nurses to a spot covered 
with damp earth, whereupon each larva is surrounded 
by a case or little dome of earth, within which it spins 
its cocoon, enwrapping the whole body. From time to 
time some worker comes to see, whether the cocoon is 
finished. As soon as it is, it is carefully cleansed of 
adhering earth, and is then stored up in a neat little 
heap in company with others that have reached the 
same maturity. With those ant-species, whose larvae 
do not spin cocoons, the larvae are not encased in earth 
before their pupation. On that account the ex- 
tremely tender skin of the pupa unprotected by a 
cocoon requires all the’*more care and caution, lest 
grains of sand or mould should enter between the 

1) On the growth of ant-eggs see Forel, “‘Fourmis de la Suisse,” 


p. 388; it is of minor importance, whether or not the increase in volume 
of ant-eggs be called growth in the proper sense of the term. 


Care of the Young in the Animal Kingdom. 165 


tiny segments of the body, or lest in transportation 
any part of the soft pupa be too sharply pinched by 
the hard mandibles of the ants, which in this case serve 
as hands. 

It requires a great deal of attention and skill on 
the part of the workers employed in nursing, only to 
keep neat and clean thousands of eggs, larvae and 
pupae. In earth-nests the moist, soft skin of these 
small beings is in continual danger of being soiled by 
sand or other foreign matter, and, besides, it is excel- 
lent soil for the growth of injurious fungi. ‘ Never- 
theless, the ants always keep their brood perfectly clean 
so that even under a magnifying lens not a speck of 
dust can be discovered. In spite of the damp and 
mouldy atmosphere, they are able to prevent entirely 
the growth of fungi both on their brood and through- 
out the nest. It might perhaps be suggested, that this 
is, for the most part, to be attributed to the antiseptic 
effect of the formic acid contained in the poison 
glands of the ants. But, in reality, the chambers in 
which the larvae are kept, show an alkaline reaction, 
as was of late pointed out by Ch. Janet,’ who explains 
this phenomenon from the fact, that the secretion of 
the epidermic, and especially of the salivary glands 
of ants is of a basic nature. 

The cleaning of the young is only a secondary 
occupation in the ant-nursery. But even in this sec- 
ondary office the ants surpass all other animals in 
care and skill. No cat by licking will wash her kittens 
with such exactness and affectionate attention, as ants 

1) “Réaction alcaline des chambres et galéries des nids de fourmis” 


(“Extr. des Comptes rendus hebdomadaires de l’Acad. des Sciences,”’ 
CXXVII, 1898, 1380). 


156 Chapter IV. 


clean the larvae entrusted to them. Therefore we are 
right in expecting, that the same perfection of the 
breeding instincts of ants will be manifested also in the 
other branches which are not less important for the 
preservation of the species, namely, in the suitable 
regulation of the temperature and in the proper nour- 
ishment and defense of the brood. 

In the care of the young it is of the utmost import- 
ance to regulate the conditions of temperature in a 
manner most advantageous to this development. In 
bee-hives the position of the brood is determined by 
the shape of the comb. Throughout its development 
the young bee remains in the same cell, in which it 
was placed as egg, and it is therefore constantly sub- 
ject to the same conditions of temperature. It is quite 
different with ants. Here the temperature has to be 
altered and regulated by the workers according to 
the different stages of development. The eggs and 
the larvae in their earliest stage are generally stored up 
in the lowest chambers of the nest, where the air is 
cool and damp. Further above the half-grown larvae 
are lodged, whilst the uppermost stories are occupied 
by the full-grown larvae and the pupae; for these 
latter require for their development greater heat, which 
is found immediately beneath the surface of the nest 
struck by the rays of the sun. If out of doors it 
‘grows chilly and rainy, the pupae and. elder larvae 
are immediately carried into the lower chambers, where 
they are better protected from cold and moisture. This 
regulation of the conditions of temperature alone, 
according to the necessities of different stages of 
development, implies astonishing sagacity, such as even 


Care of the Young in the Animal Kingdom. 157 


men would be able to acquire only after years of 
observation and study. Yet, in ants every single worker 
is endowed with it as soon as she is drawn from her 
cocoon and has become dry. This is because their 
sagacity is instinctive, essentially different from intel- 
ligence and reflection. Ants are in their every action 
guided directly by sensitive perceptions, not by intel- 
lectual ideas. The enigma, therefore, is satisfactorily 
explained by the innate adaptation of their sensitive 
cognition and appetite, whereas the hypothesis of ani- 
mal intelligence is unable to offer any solution. 

But now we come to the most puzzling and mys- 
terious question in the nursing of ants, namely, the 
influence of the education of the young larvae on the 
development of different castes in ant-states. Science 
has but just now begun to divine the mysteries hidden 
here; but it is still far from having fathomed their 
depths. We shall very briefly place before our readers 
what is certain or at least probable concerning this 
matter. It will fully suffice to prove, that the nursing 
instincts of ants, bees and termites are far superior 
to those of birds and mammals. 

According tothe older opinion, thus far commonly 
held, and based chiefly on Dzierzon’s classical obser- 
vations on bees, the sex of their posterity is determined 
by the instinctive choice of the oviparous queen, and 
not by the workers that rear the brood. Because it 
has been observed, that with ants, bees and wasps 
unfertilized eggs produce males only, it is assumed 
that also the normal males of these social insects are 
always hatched from unfertilized eggs. The queen, 
when depositing her eggs, is supposed, by either open- 


158 : Chapter IV. : 


ing or closing the connection between the oviduct and 
the seminal vessel, to control, under the influence of 
her oviparous instinct, the fertilization of the egg, and 
therefore to decide, whether it would develop into a 
male or a female. This instinct of the queen is aroused 
to suitable activity by the peculiar nature of the cell, 
into which she puts her head before oviposition; in 
drone-cells she puts an unfertilized egg, in those of 
a future queen or worker a fertilized one. Probably 
she is led to make this difference not so much by the 
touch-perception of the different shapes of the cells, 
but rather by the smell of the salivary gland secretions 
employed by the workers in their construction.t The 
peculiar odor of the cells, however, as well as their 
size and form is due to the architecture of the workers ; 
thus, in bees the sex of the offspring is indirectly 
at least controlled by the instincts of the workers. 
With ants it is different, because their queens do not 
deposit the eggs in cells, but simply suffer them to 
be received and carried away by the workers. The 
instinctive self-determination of the oviparous female 
seems, therefore, to be greater with ants. Of course, 
it is very probable, that the nourishment and treat- 
ment of the queen on the part of the workers indirectly 
also influences the oviposition; but in what manner, 
is as yet entirely unknown. 

To pass from the queen to the workers, it was ascer- 
tained long ago, that, with social wasps, bees and ants, 
also workers, which have small ovaries with a reduced 
number of ovarial tubes, and are, besides, unable to 

1) Cf. on this point N. Ludwig, ‘‘Futtersaft oder thierische Veran- 


lagung,” p. 32; and p. 57 of the publication of Ferd. Dickel mentioned 
below. 


Care of the Young in the Animal Kingdom. 159 


mate, nevertheless sometimes lay eggs capable of 
development. This :phenomenon has been called by 
Ch. v. Siebold,t to whom we are specially indebted 
for its discovery, virgin-generation, or parthenogene- 
sis. Under natural conditions parthenogenesis occurs 
with ants principally in colonies which have lost their 
queen, and therefore try to rear their posterity from 
eggs laid by workers. In several observation-nests 
of Polyergus rufescens, Formica sanguinea and rufi- 
barbis the queen was missing, and I observed that the 
workers,—with Polyergus the slaves,—selected an extra 
large worker of the dominant species as a substitute 
for the queen, treated her with greater care, gave her 
more food, and thus induced her to parthenogenetic 
oviposition. This shows that under certain circum- 
stances the instinct of ants is able to effect by special 
treatment the development of the ovaries of even 
adult workers, so as to make them capable of laying 
eggs ;? but these unfertilized eggs can only produce 


1) “Wahre Parthenogenese bei Schmetterlingen und _ Bienen,” 
Leipzig, 1856. Quite recently Ch. Janet has discovered parthenogenesis 
also among hornets (Sur Vespa Crabro, Extr. des Mém. de la Soc. 
Zool. de France, 1895, p. 75). 

2) This form of parthenogenesis, which is spontaneouly caused by 
the workers themselves, must be carefully distinguished from another 
form caused by artificially raising the temperature, and quite independent 
of the ants’ instinct. See my observations in “Biolog. Centralbl.,” XI 
(1891), No. 1: “Parthenogenesis bei Ameisen durch kuenstliche Tem- 
peraturverhaeltnisse.” The experiments made by E. Bickford (“Ueber 
die Morphol. und Physiol. der Ovarien der Ameisen-Arbeiterinnen,” 
in “Zool. Jahrb. Abth. fuer Systemat.,” IX, 1895, 1st issue) with 
Lasius fuliginosus (p. 19; Sep., p. 23) belong rather to the second 
category than to the first, since she too employed artificially raised 
temperature. At any rate, they do not approach natural conditions as 
closely as my observations mentioned above. On the latter cf. ‘‘Stett. 
Entom Ztg.,’’ 1890, pp. 303-305, and ‘‘Biolog. Centralbl.,” 1895, pp. 
609 and 610. 


160 ’ Chapter IV. 


males.t. This phenomenon manifests the marvelous 
sagacity and quasi-intelligent plasticity of animal 
instinct, which can hardly be styled “automatism.” 
Neither can it be identified with intelligence properly 
so-called, for this would suppose rational knowledge 
of the internal laws governing the growth of the ant- 
organism, a knowledge far surpassing even the intelli- 
gence of man and entirely beyond the reflections and 
experience of ants. Only the appropriate disposition 
of their sensitive cognition and appetite can account 
for the fact, that the perception of a given want is 
followed by a corresponding modification in their 
nursing instinct, by which the defect in question is 
remedied. 

According to Dzierzon’s views, which we men- 
tioned above, it is the oviparous instinct of the queens, 
that controls the sex of the bee developed from a 
given egg; in this supposition the worker-bees are 
assigned a merely indirect influence. Of late, how- 
ever, another theory on the differentiation of castes 
in bees has been advanced, which assigns to the nurs- 
ing instincts of the workers a far more extensive 
sphere of action. The originator was an Italian priest, 
Lanfranchi by name, who published it in 1894 in the 
“Apicoltore.” In Germany it was developed and 
confirmed by new experiments, principally by Ferd. 
Dickel,? the editor of the “Noerdlinger Bienenzeitung.” 


1) More recently H. Reichenbach has published some observations 
(in “Biol. Centralbl.,” 1892, p. 461 ff.) which seem to prove, that with 
Lasius niger the parthenogenetic eggs laid by workers may give origin 
also to workers. But further confirmation will be required before 
accepting this statement. In North America Prof. W. M. Wheeler has 
lately published some interesting reports on parthenogenesis in ants. 

2) “Das Princip der Geschlechtsbildung bei Thieren geschlechtlicher 
Fortpflanzung, entwickelt auf Grundlage meiner Bienenforschungen.” 
Noerdlingen, 1898; cf. especially p. 20. 


Care of the Young in the Animal Kingdom. 161 


Dickel says: “Under normal circumstances the fer- 
tilized mother-bee lays only fertilized eggs; it is the 
workers, that influence and control the fate of these 
homogeneous eggs.” According to this new opinion, 
also those eggs, which in normal bee-hives produce 
drones, are fertilized; and not only the differentiation 
between queen and worker, but between queen and 
drone, and between worker and drone, is due to 
the influence of the salivary gland secretions of the 
workers on the eggs previously deposited in the cells. 
Hence, Dickel regards both queens and drones only 
as the foundation for the development of the sexes, the 
workers, however, as the really determining factors. 
According to Dickel, certain salivary glands of the 
workers contain the secretions determining the sex, 
and the differentiation of all the castes in bee-hives 
depends on the instinctive application of these secre- 
tions, when the workers are licking the eggs. 
Although several biological experiments of other 
authors seemed to confirm the theory that under nor- 
mal conditions all the eggs in a bee-hive are fertilized,’ 
we must add, nevertheless, that the very exact micro- 
scopical studies of Paulke and Petrunkewitsch on the 
existence or non-existence of spermatozoids in the 
eggs of bees rather corroborate the old theory of 
Dzierzon, according to which the eggs giving origin 
to drones develop without containing any spermato- 


- 1) Cf. N. Ludwig, ‘Neues ueber Ernaehrungs- und insbesondere ueber 
Fortpflanzungsverhaeltnisse der Honigbiene” (‘‘Natur und Offenb.,” 
XLIV, 1898, 12th issue, pp. 705-719), ““Weiteres zur neuen Lehre ueber 
die Geschlechtsbesti g der Bienen” (ibid. XLV, 1899, 3d issue, pp. 
140-148; “Weitere Ergebnisse ueber die Fortpfl gsverhaeltnisse der 
Biene” (‘‘Natur und -Offenbarung,” 1901, 7th issue, pp. 426-430). 

it 








162 Chapter IV. 


zoids. It is best, therefore, to suspend our judgment 
on this problem, until it is definitely solved. 

If the new theory of the fertilization of all the eggs 
in a bee-hive, under normal conditions, should prove 
true, it ought to be extended also to the ants. Hence, 
in ant-colonies, too, it would be the workers, who by 
their nursing instincts determine, whether a given ant- 
egg is to produce a worker, a winged female, a soldier, 
or a male. 

We intend here, however, to consider only the 
wonderful influence exercised by nursing on the dif- 
ferentiation between females and workers. This is an 
established fact, quite independent of the new theory. 
Why is it that one and the same kind of egg now 
produces a queen with complete power of generation, 
now a worker devoid of generating powers, but com- 
pensated, as it were, for this loss by psychic endow- 
ments all the more perfect? Here we enter a mys- 
terious region, where the breeding instincts of social 
insects reign supreme, an instinct which for its creative 
power is unparalleled in the entire animal kingdom. 

It is a well-known fact, that with honey-bees a 
worker larva can be developed into a queen by increas- 


1) W. Paulke, in ‘“‘Anatomischer Anzeiger,” Vol. XVI, 1899; A. 
Petrunkewitsch, “Tie Richtungskérper und ibr Schicksal im befruchteten 
und unbefruchteten Bienenei,’” (Zool. Jahrbuecher,” Abtl. fuer 
Anatomie, Vol. XIV, 1901); Aug. Weismann, “Ueber die Partheno- 
genese der Bienen” (“‘Anatom. Anzeiger,” Vol. XVIII, 1901, Nos. 20-21); 
H. v. Buttel-Reepen, ‘‘Ueber die Dzierzon’sche Theorie” (‘‘Bienen- 
wirtschftl. Centralbl.,”’ 1901, No. 1); “‘Der Abschluss der Freiburger 
Eiuntersuchungen” (Ibid., 1901, No. 19); “Die Parthenogenesis bei 
der Honigbiene” (“Natur und Schule,” Vol.I, 1902, 4th issue); P. 
Bachmetiew, “Ein Versuch, die Frage ueber die Parthenogenese der 
Drohnen zu lesen” (“‘Allgem. Zeitschr. f. Entom.,”’ 1903, Nos. 2-3). 


Care of the Young in the Animal Kingdom. 163 


ing its cell and giving it different food.t Also in the 
case of termites, so we are informed by Grassi and 
Sandias,? the various methods of nursing the larvae, 
and especially the different salivary gland secretions of 
the nurses are of great importance in the differentia- 
tion of the castes of one and the same species. The 
same probably holds good with ants,* and, indeed, with 
far more variability than with honey-bees, although less 
than with termites. This is indicated by the numerous 
intermediate forms between the females and workers 
of ants. Of course, wherever these appear, they are 
exceptional forms, yet none the less they throw some 
light on the origin of the normal differentiation into 
females and workers; for, their existence is most 
intelligible on the supposition, that the difference of 
caste is not predetermined in the egg, but that it will 
depend on the nursing, whether the fertilized egg will 
bring forth a winged, perfect female, a normal worker 
or perhaps some intermediate form. 

Of course, the specific development peculiar to 
every ant-species is the necessary foundation for the 
differentiation of the normal castes and for the origin 
of certain abnormal, intermediate forms. Where there 








1) Cf. N. Ludwig, “‘Futtersaft oder thierische Veranlagung.”— 
According to Planta’s tables the food of queen-bees contains a far 
larger amount of fat. But according to Ludwig it is especially the 
different quality of the saliva, added by the bees to the nutrifying 
juice, which is of decisive importance. 

2) “Costituzione ¢ sviluppo della Societa dei Termitidi” (Catania, 
1893), pp. 75-106. 

8) Emery, ““Le Polymorphisme des fourmis et la castration alimen- 
taire,’’ Leyden, 1896 (Extr. du Compte rendu des Séances du 3me 
Congrés internat. de Zool., p. 395 ff.). 

*) Wasmann, “Die ergatogynen Formen bei den Ameisen und ihre 
Erklaerung” (‘‘Biolog. Centralbl.,’’ 1895, Nos. 16 and 17). 


164 Chapter IV. 


is no possibility of developing into certain forms, there 
is, of course, no basis, no material for the exercise of 
the nursing instincts of the workers. From this it 
becomes clear, why fixed intermediate forms between 
females and workers occur with certain species, dif- 
ferent forms with other species, whilst with others 
again there are none at all. But, within the limits of 
this natural disposition for further development, there 
remains to the ants a wide range for exercising their 
nursing instincts. 

These intermediate forms between females and 
worker ants I have grouped into six classes, but here 
we are concerned with but one or two of them. In 
some of these “ergatogyne” forms it appears almost 
at a glance, how they came into existence, namely, 
whether the larva, reared up to a certain stage to be 
a worker, was later on cared for so as to become a 
female, or whether the opposite took place. In the 
first case the intermediate form makes the impression, 
that the worker-character had been developed to 
excess; in the second case, that the female character 
had been stunted; in the former the so-called worker- 
like (ergatoid) queens are the result, in the latter a 
kind of female-like workers, which I have named 
pseudo-females (pseudogynes). The former combine 
the vaulted thorax of females with the small and 
abdominal development of queens; the latter unite 
the vaulted thorax of females with the small and 
stunted abdomens of workers. Especially the latter 
form, the pseudogynes, are apparently best accounted 
for on the score of education rather than by a peculiar 
disposition inherent in the egg, from which they are 


Care of the Young in the Animal Kingdom. 165 


hatched.1 I know some colonies of Formica san- 
guinea, near Exaten, in which these pseudogynes sud- 
denly made their appearance, became more numerous 
in the course of the next few years, and gradually 
decreased later on, or disappeared entirely. One 
colony (No, 21) in 1895 reared every possible pseu- 
dogyne intermediate form between normal workers 
and normal queens! Since the queens which lay the 
eggs in these nests are unable to change at will the 
nature of their ovaries from year to year, but are 
always compelled to lay fertilized eggs, capable of 
equal development, the origin of those intermediate 
forms is probably due to changes in the manner of 
nursing, and to modifications in the very nursing 
instincts of the workers. This supposition is con- 
firmed by the fact, that in F. sanguinea there is a 
certain causal relation between the origin of pseu- 
-dogynes and the education of the larvae of a genuine 
ant-guest, the beetle Lomechusa strumosa. I have 
ascertained this mysterious connection by means of 
my statistics embracing 410 sanguinea colonies within 
a radius of several kilometers around Exaten; these 
statistics will be published later on in some scientific 
periodical.? Here it may suffice to mention, that the 
centres of propagation of the pseudogyne forms and 
of the Lomechusas are always together in the same 

1) See my recent publication, ““Neue Bestaetigungen der Lomechusa- 
Pseudogynen Theorie’ (‘“Verhandl. der Deutsch. Zool. Gesellsch.,’’ 1902, 
pp. 98-108 and Pl. II), where this theory is extended also to North 
American ants. See below (the following section, p. 179 foll. and the 
plate opposite p. 181). 

2) The beginning of those statistics dates back to 1895 (“‘Die 


ergatogynen Formen bei den Ameisen und ihre Erklaerung,” “Biol. 
Centralbl.,’”? 1895, Nos. 16 and 17). 


166 Chapter IV. 


or at least in neighboring nests. The number of 
colonies, in which I found Lomechusas (100), is more 
than ,three times as great as that of nests containing 
pseudogynes (33); these are the centers, from which 
the Lomechusas gradually spread to the neighboring 
nests, where by and by they cause the birth of pseu- 
dogynes. It is scarcely possible, that the presence of 
these beetles and of their larvae, which are fed by the 
workers, should have a modifying influence on the 
ovaries of the queens, but, probably they do so on 
the nursing instincts of the workers. My recent 
observations and experiments until 1904 have con- 
firmed this solution of the interesting problem. 
Some of the above mentioned intermediate ant- 
forms are useful for the preservation of the colony 
and the species, whilst others are more or less indif- 
ferent, and still others positively hurtful, being prob- 
ably pathological deformities. The rearing of worker- 
like, wingless queens among the Amazon ants (Polyer- 
gus rufescens), for instance, is very appropriate, 
because their colonies are rather rare and far distant 
from one another. Therefore, the probability is very 
slight, that on their nuptial flight the winged sexes 
will meet with those of other colonies; and besides, 
this species has to encounter exceptional difficulties in 
founding new settlements by means of single fertilized 
females, since the Amazons are entirely dependent on 
the help of their slaves. The wingless queens, on the 
other hand, cannot go far from their nests; and after 
they have been impregnated by the winged males, 
some strolling slaves can easily find them in the 
neighborhood and bring them home again; hence it 


Care of the Young in the Animal Kingdom. 167 


is very suitable, that just Polyergus should so often 
rear ergatoid queens. Nor is this arrangement in 
any way due to the “intelligence” of the Amazons; for 
the education of their offspring is entirely committed 
to the care of their slaves (mostly F. fusca) ;+ these 
slaves, however, have been robbed from colonies which 
do not rear ergatoid females, and neither reflection 
nor experience could have given them intellectual 
knowledge of the requirements peculiar to the nursing 
of Polyergus. Here animal intelligence is entirely 
powerless. If F. fusca, the slaves of the Amazons, rear 
the offspring of their masters in a way suited to the 
preservation of exactly this species, then we must 
admit, that the nursing instincts of the slaves are 
influenced and modified by the peculiar sensations 
caused by the Polyergus-nests. 

But what shall we say of the rearing of pseu- 
dogynes with F. sanguinea? This combination of 
female and worker is decidedly injurious to the 
preservation both of the colonies and of the species. 
The pseudogynes are stunted beings, neither workers 
nor females, unfit to participate either in building the 
nest or in nursing the young,’ in defending the colony 
or in propagating the race; in fine, they are down- 
right failures. It is evident, that their origin is not 
due to the “individual intelligence of the ants”; for, 


1) Near Exaten, Holland, all Polyergus-nests, I met with, contained 
F. fusca as slaves; those which I found in Bohemia (Mariaschein), 
Austria (Vienna) and in Luxemburg, had F. rufibarbis as slave species. 
The ergatoid queens I met hitherto only in nests with fusca-slaves. 

2) It happened only very seldom (among several hundred observa- 
tions only five times), that, on the nest being exposed to the light, a 
pseudogyne seized and carried away an ant-larva, whilst workers are 
always wont to do so, 


168 Chapter IV. 


if they had but a spark of intelligence, sad experiences 
would have enlightened them long ago on the folly 
of this mistake. Nay more, if the pseudogynes owed 
their origin to the normal nursing instincts of ants, 
we should have to despair of the fitness of animal 
instinct and even of the wisdom of the Creator. What 
is the key to this mystery? It is the beetle Lomechusa 
strumosa. According to our hypothesis the rearing 
of the pseudogynes is an aberration of the breeding 
instinct of ants, caused by the continuous education 
of Lomechusa-larvae. In the economy of nature it 
is the duty of this beetle, to check the excessive 
increase of the ant-species, whose hospitality it enjoys. 
For this reason its larvae not only consume countless 
ant-eggs and ant-larvae,—the ants calmly looking on 
‘the while,—but by destroying the offspring of the 
ants, and by the care which the ants bestow on them, 
they cause the degeneration of the normal nursing 
instincts of the workers, resulting in the education of 
crippled pseudogynes.t. To account for these facts on 
the score of “individual animal intelligence” would 


1) These expositions will probably suffice also to refute an objection 
raised by Dr. G. Adlerz, who, misunderstanding my psychological ex- 
planation of the rearing of pseudogynes, says in the third part of -his 
valuable “Myrmecologiska studier’ (Stockholm, 1896), p. 51: ‘With 
regard to this Wasmann seems inclined to credit ants with an exag- 
gerated power of reflection, which he otherwise is unwilling to do.” 
Besides, the pathological degeneration of the breeding instinct explains, 
why the rearing of pseudogynes is still continued, even when colonies 
have been deprived of their Lomechusas. By the way, let me repeat 
a remark formerly made, that the causal connection of pseudogynes with 
the Lomechusas is not to be confounded with the explanation of this 
connection. The former seems to be firmly established by direct observa- 
tion, the latter is still an hypothesis, but an hypothesis, strongly con- 
firmed by recent experiments of myself and of Vichmeyer. See ‘Neue 
- Bestaetigungen der Lomechusa-Pseudogynentheorie” (‘‘Verhandlungen 
der Deutsch. Zool. Gesellsch.,’’ 1902, p. 98 ff.). 


Care of the Young in the Animal Kingdom. 169 


involve us in endless and hopeless contradictions. 
They are explainable only from the standpoint of a 
higher, teleological consideration of nature, which 
does not presume to replace the wisdom of the Creator 
by the “intelligence of animals.” 

The phenomena in the nursing of ants mentioned 
above can be ascertained only by close, scientific 
observation. But some other features are known to 
every amateur in the study of ants, and do not escape 
even the most casual observer. The first thing that 
strikes our attention is the great attachment displayed 
by the. workers for their charges. They carefully 
guard them against every disturbance, and at the risk 
of their own lives they exert all their strength in ward- 
ing off hostile invaders. The whole colony is seized 
with frenzy, if an attempt is made to rob them of 
their larvae and pupae.t. Merely thrust your stick 
into a hillock of wood-ants! At once there ensues a 
tumultuous uproar and masses of workers rush forth 
to rout the enemy. But if you happen upon a chamber 
filled with pupae and attempt to take away the cocoons, 
the fury of the ants reaches its climax. Like an army 
of raging furies they fall upon the assailant, viciously 
biting and ejecting their poison. Hundreds and thou- 
sands are crushed by the enemy, but other hundreds 
and thousands are eager to face the carnage. No 
lioness, no she-monkey ever defends her young with 
the heroism displayed by ants. Workers will rather 

1) This is the case with species otherwise very peace-lovirig, e. g., 
with the large American leaf-cutting ants of the genus Atta. On open- 
ing a nest of Atta sexdens in Rio Frio, Forel even had an artery of his 
little finger pierced by a large-headed worker. See Forel, “Zur Fauna 


und Lebensweise der Ameisen im Columbischen Urwald” (“Mittheil. der 
Schweiz. Entom. Gesellsch.,”? Vol. IX, 9th issue, p. 407). 


170 Chapter IV. 


suffer their heads to be torn off, than yield to the 
enemy the pupae they carry in their jaws. And yet, 
it is not even for their own children, that they ‘‘sacri- 
fice themselves so unselfishly ;’ their charges are but 
their foster-children. But that higher natural law, 
which has made preservation of the species the fore- 
most instinctive commandment implanted in the ani- 
mal soul, this natural law, I say, also constrains the 
worker-ants to risk life and limb in behalf of beings 
begotten by others. This commandment they observe 
faithfully, not led by any sense of duty or by noble 
forgetfulness of self, but by an irresistible, instinctive 
impulse implanted in them by Another, and to which 
they yield obedience, not intelligently or voluntarily, 
but urged on by a blind necessity of nature! 

To credit animals with intelligence, to ascribe to 
them ever so faint a trace of intellectual knowledge of 
the purpose of their actions, will necessarily lead to 
extolling the self-sacrifice of the single workers for 
the welfare of the colony and especially for the young, 
as a high degree of quasi-human, nay of superhuman 
virtue. And in fact, L. Buechner, E. Haeckel, Th. 
Eimer, O. Zacharias and other modern animal psychol- 
ogists have actually ventured such assertions.1 Of 
course, their only commendation is their boldness, but 
it is a boldness leading to the greatest absurdities. 

What is it then, that impels the ants to such heroic 
devotedness and self-sacrifice for the offspring of 
their colony? Is it perhaps “motherly love’? No; 
for the workers are but the sisters or aunts of their 


1) Wasmann, “Die zusammengesetzten Nester und gemischten 
Kolonien der Ameisen,” pp. 190 and 191, 


Care of the Young in the Animal Kingdom. 171 


charges, since under ordinary circumstances the eggs 
are laid only by the impregnated females. Their 
affection, therefore, as sisters or aunts is the psychic 
impulse of their nursing instinct. According to 
modern animal psychology, which ascribes to animals 
besides their instinct at least a modicum of genuine 
intelligence, it cannot be doubted, that the worker- 
ants fulfill their duties as sisters or aunts “very know- 
ingly,” that they are aware of the importance of the 
work allotted to them in their social economy, and 
that they apply themselves to nursing the offspring 
of another with the consciousness of doing their duty. 
But to what degree will the love of the workers for 
the young be advanced, if to their affection as sister 
or aunt there is added “motherly love’ in the full 
sense of the term? Must we not expect that the love 
of the workers for their own young should attain to 
an exalted, an unutterable degree of tenderness? For, 
in the whole creation no love is stronger than that of 
a mother. 

Indeed, our expectations would be justified, if ants 
were endowed with intelligence and self-consciousness. 
But what do we find in reality? The workers gen- 
erally devour most of the eggs, which they have laid 
themselves.1. Is this the climax of noble, self-sacri- 
ficing motherly love? Or shall we call these workers 
abominable cannibals regardless of duty? The psy- 
chology of Brehm and others of that ilk may decide 
this question. In our opinion, however, such facts 
ought to lead reasonable people to perceive the obvious 
contradictions, in which all the talk about “animal 


1) See my observations-in “Biolog. Centralbl.,” XI, 1891, p. 21 ff. 


172 Chapter IV. 


intelligence” and ‘animal ethics’ is hopelessly 
involved, 

The nursing instinct of ants with all its “devoted- 
ness and unselfishness,” is therefore nothing else than 
a purely instinctive impulse guided and determined 
in its operations only by sensitive impressions and not 
by intellectual concepts. Under normal circumstances 
this instinctive impulse is appropriately regulated, and 
manifests itself as the product of “self-sacrificing 
sisterly love.” But, if the abnormal irritation of the 
nervous system of the ants, caused by parthenogenesis, 
has disturbed the normal sphere of sensitive impres- 
sions, then sisterly love is not changed into motherly 
love, but into “barbarous, unfeeling cannibalism”’! 

Modern animal psychology evidently toys in a 
rather frivolous manner with the term “motherly 
love,” by applying it to the nursing instincts found 
among animals. Nor can the plea be advanced, that 
with higher animals matters are quite different than 
with ants; for, we have proved above, that the nurs- 
ing instincts of ants far surpass in perfection those of 
birds and mammals, not only by their quasi-intelligent 
self-determination in the method of education, but 
also by the great unselfishness manifested in nourish- 
ing and defending their young. If there should be 
any difference at all, it is in this, that in the care of 
their offspring the higher animals betray far less 
“intelligence” and far less “individual liberty,” than 
is found in ants. Moreover, it is a well-known fact, 
that domestic pigs not seldom devour some of their 
litter; yet pigs are “higher animals.” In such cases, 
however, the sow sins as little against good morals, 


Care of the Young in the Animal Kingdom. 178 


as worker-ants do by devouring their own eggs; for, 
morality presupposes reason and free-will, reflection 
and consciousness of duty, all of which are wanting 
throughout the animal kingdom, being the exclusive 
privilege of man. 

That animals in caring for their young are not led 
by reason, but only by sensitive emotions and repre- 
sentations, becomes evident especially from the phe- 
nomena of adoption in the animal kingdom. There- 
fore these shall form the subject of the following 
section. 


3. Adoption Instincts in the Animal Kingdom. 


The tendency to adopt the offspring of strangers 
is shown by all those animals which, to preserve their 
species, are forced to bestow great care on their own 
progeny. This tendency is found among ants not 
only with regard to the eggs, larvae and pupae of 
other colonies of their own species or of allied species, 
but also with regard to members of altogether dif- 
ferent orders of insects, living in their communities. 
These adoption instincts are responsible for the mixed 
colonies of slave-making ants, the robbed pupae of the 
slave-species being nursed as carefully as others, either 
by the masters or by the slaves already present in the 
nest. To the same instinct of adoption must be 
referred the care bestowed by the ants on their gen- 
uine guests or other nest-mates belonging to different 
orders of insects, but above all, the solicitude 
with which they rear the larvae of certain beetles 
(Lomechusa, Atemeles, Xenodusa) and the eggs of 
several kinds of plantlice. The adjoining illustration 


174 Chapter IV. 


shows the beetle Lomechusa strumosa so often referred 
to, and one of its larvae, magnified to four times their 
natural size. 





Fig. 4. Fig. 5. 
Lomechusa strumosa F. Fullgrown larva of Lomechusa 
(Magnified. ) strumosa. (Magnified.) 


The same adoption instinct occurs also with birds, 
although not so seemingly intelligent as with ants. 
The best-known example is the hen, that readily 
hatches eggs of other hens, ducks, geese, turkeys, 
etc., and extends to all her adopted children the same 
“motherly care,” she would show to her own chicks. 
G. Romanes? succeeded even in making a hen the 
foster-mother of some young ferrets, which he had 
substituted for the artificial eggs, on which she was 
hatching. The numerous species of birds, which 
tend the young cuckoos, follow the same line of con- 
duct, the only difference being, that they lavish still 
greater care on these changelings, because they open 
their mouths wider in crying for food than their own 
nestlings. The adoption instinct, finally, is met with 
among mammals, the most blood-thirsty carnivores not 
excepted. Though it is a fable, that ancient Rome 


1) “Mental Development in the Animal Kingdom.” 


Care of the Young in the Animal Kingdom. 175 


owed its origin to the adoption instinct of a she-wolf 
that gave suck to Romulus and Remus, yet similar 
facts are fully authenticated; for instance, that suck- 
ling cats adopt young rabbits and squirrels.1 Espe- 
cially among apes this instinct is quite prominent, but 
much more characteristically unreasonable than with 
any other animal. Of course, for a certain kind of 
modern animal-psychology, which is accustomed to 
the most superficial observation, such occurrences are 
noble manifestations of human compassion, and of an 
abundance of “motherly tenderness” lavished on the 
offspring of strangers. But accurate, scientific obser- 
vation and critical investigation prove the very con- 
trary, namely, that throughout the animal kingdom 
the nursing instinct is but a sensual impulse, unac- 
companied by individual intelligence or individual 
morality. 

If a hen calmly continues trying to hatch pieces of 
limestone or links of iron chains put in place of her 
eggs, she can hardly be said to be actuated by 
“motherly love.” Animals merely endeavor to satisfy 
their instinctive breeding impulse; the higher pur- 
poses of their action are altogether unknown to them. 
William of Reichenau relates that a bitch, being rob- 
bed of her pups, fetched an old pair of slippers and 
tried to suckle them.?, Whether she thereby intended 
to allay her pangs of conscience as to the fulfillment 
of her “maternal duties,” animal psychologists a la 
Brehm will be better able to decide than we, to whom 
such facts merely prove, that the nursing instinct in 


1) Cf. W. Herd, in “Scottish Naturalist,” 1872, p. 155. 
2) Cf. “Kosmos,” 4th year, VII (1880), p. 217, 


176 : Chapter IV. 


animals is a sensual impulse, not guided by reason 
and reflection. 

This organico-sensitive nature of the nursing 
instinct also explains, why-it often extends to the 
helpless offspring of other species, whose instinctive 
behavior is somewhat similar to that of the animal’s 
own progeny. The sense-perception of these helpless 
beings stimulates the nursing instinct of the old ones, 
and therefore they “adopt” the young. ones of 
strangers. The smell of the larvae of Lomechusa 
strumosa is especially attractive to the sanguine slave- 
makers; besides these larvae instinctively mimic the 
attitudes and behavior of the ant-larvae, and although 
they possess six feet, they do not make use of them, 
but conduct themselves like helpless ant-larvae. For 
these reasons they enjoy the most careful attention on 
the part of their hosts. And as these beetle-larvae, 
when fed by the ants, grow much faster than the ant- 
larvae, they impress the instinctive nursing impulse of 
the ants far more favorably than the latter, and hence 
are the objects of “greater tenderness.” At any dis- 
turbance of the nest the workers first care for their 
“adopted children” and bring them to a place of safety, 
before they attend to their own offspring; yea, they 
even neglect the rearing of the latter, their only care 


1) That the ants do not confound those coleopterous larvae with 
their own on account of their shape and color, I ascertained, in May, 
1897, by experiments with larvae of Anthonomus pomorum, which are 
far more similar to ant-larvae than those of Lomechusa. The Antho- 
nomus-larvae were instantly seized as prey and torn to pieces by the 
sanguineas of my observation nest. On the whole, it must not be 
imagined that the Lomechusa-larvae make the same impression as their 
own on the sensitive perception of the ants; the impression is at most 
similar, but more agreeable, which probably explains why the ants prefer 
the adopted larvae to their own. 


Care of the Young in the Animal Kingdom. 177 


being their Lomechusa-larvae, so dear to them on 
account of their quicker growth and their better 
appetite. It does not affect the ants in the least, that 
the Lomechusa-larvae again and again devour the 
eggs and young larvae of the ant colony by the whole- 
sale; on the contrary, they even carry these change- 
lings to the clumps of eggs and larvae to facilitate 
their work of destruction. Unless a superior Wisdom 
had provided that the ants themselves, by their stupid 
affection, prevent the excessive increase of the Lome- 
chusa population, the number of these guests would 
become so large as to destroy all the sanguinea- 
colonies. But there is no danger of any such calamity ; 
for, the ants deal with the Lomechusa-larvae during 
their pupation just as they do with their own, imbed- 
ding them carefully in a vault of earth. After a 
short time, the ant-larvae having meanwhile spun 
their cocoons, are again removed from the earth. 
This latter measure applied to the Lomechusa-larvae 
proves fatal to them. The larvae of these beetles do ° 
not spin a solid cocoon, but only an extremely flimsy, 
silken web, which tears as soon as they are unearthed ; 
soon after the Lomechusa-larvae are again carefully 
imbedded at some other place, then they are taken 
out again, carried about, again imbedded, until at 
length they become dry and perish. In this manner 
the folly of the ants causes most of the Lomechusa- 
larvae to die before they are changed into pupae; and 
even those, which have fortunately entered the state of 
a pupa, are often unearthed by the ants and devoured, . 
—perhaps from an excess of affection? According 
to my long continued observations on the development 
12 


178 Chapter IV. 


of Lomechusa strumosa in normal sanguinea-colonies 
only those larvae escape destruction, which, after hav- 
ing been imbedded in their little cave, are forgotten 
by the ants; all the rest are hopelessly doomed; of 
100 larvae, therefore, at most about Io reach the stage 
of imago, sometimes scarcely one. For instance in 
the observation nest illustrated on p. 23 in May, 1896, 
about 150 Lomechusa-larvae, the offspring of Io 
Lomechusas, were reared under the most favorable 
conditions both of nutrition and temperature; from 
these 150 larvae I obtained—one single Lomechusa! 
For thousands of years and in thousands of normal 
colonies F. sanguinea year after year repeat the same 
senseless performance: first, with the greatest 
devotedness they nurse the Lomechusa-larvae, even 
allowing their own offspring to be devoured by them; 
then, their stupid affection does not allow them to 
leave the larvae in peace during the time of pupation, 
and finally they devour the pupae. They cannot see 
that during their pupation Lomechusa-larvae are to 
be treated differently from those of ants; but this is 
their salvation; for otherwise their care of the Lome- 
chusas would long ago have brought about the ruin 
of their own race. One and the same superior Wis- 
dom has designed, that on the one hand the increase 
of the ants be checked by their inconsiderate love for 
Lomechusa strumosa and for their larvae, and that on 
the other hand the spread of this beetle be kept within 
limits by the very same unreasonable affection of the 
ants. By these means so gentle and yet so effective, 
Divine Wisdom is able to maintain the equilibrium in 
nature. In the face of such phenomena, the defenders 


Care of the Young in the Animal Kingdom. 179 


of animal reason and animal ethics stand utterly 
bewildered. 

We have seen so far, how the Lomechusa-larvae 
are treated in normal sanguinea-colonies, in which the 
education of Lomechusas dates but one or two years 
back. According to my observations during the last 
years, the case is, however, different in colonies, where 
the influence of the parasitic larvae of that beetle has 
caused the appearance of those crippled, intermediate 
forms between females and workers, which we have 
called pseudogynes. For the reason mentioned above, 
only a few Lomechusas are developed, as a rule, in 
the normal colonies, whereas in colonies containing 
pseudogynes, many more of the beetles pass their 
pupation successfully, because the pupae generally 
remain undisturbed. This accounts for the fact, that 
the sanguinea-colonies containing pseudogynes are the 
centers from which the rapidly multiplying Lomechusa 
infests also the neighboring nests. One instance may 
be mentioned here. Colony No. 191 of my statistical 
map in the middle of May, 1898, had from 2 to 3% 
of pseudogynes; in August, however, the number of 
pseudogynes newly developed during summer had 
risen to 30%! In order to ascertain, how many beetles 
had been reared in this colony in 1898, I dug up the 
nest at the end of September and found among the 
ants 116 Lomechusas' snugly ensconced in their winter- 


1) From this number the 30 Lomechusas were taken, which I put 
in my sanguinea nest at home and the greater part of which were again 
driven out by the ants (see p. 59). This surprising conduct is some- 
what accounted for by the fact that the colony of my observation nest 
belonged to the normally developed class, and not to such as contained 
pseudogynes, 


180 Chapter IV. 


quarters. The rapid increase of pseudogynes in that 
colony, therefore, was in proportion to the number of 
beetle-larvae which had successfully developed in the 
same nest that year. 

It is, therefore, necessary to assume, that in san- 
guinea colonies the frequent rearing of Lomechusa- 
larvae gradually modifies the normal nursing instinct 
of the ants. This modification is manifested partly by 
the production of the crippled pseudogynes, partly by 
the more appropriate treatment of the Lomechusa- 
larvae which, after having been imbedded in their 
cradles, remain undisturbed. Ants, therefore, grad- 
ually learn to modify their nursing instinct. Is not 
this a proof of intelligence? True, their sensitive cog- 
nition guiding their instinctive activities may furnish 
the immediate occasion for that two-fold modification. 
But we have proved in a former essay, in discussing 
the different forms of learning,t that not every modi- 
fication of the hereditary instinct, occasioned by sense- 
experiences, is due to intelligence, but only that, which 
manifests a knowledge of the appropriateness of a 
given action. If ants were gifted with intelligence, 
they could not help understanding, that by improving 
their treatment of the Lomechusa-larvae, they cause 
their colony but to perish the sooner, just as they con- 
demn it to utter destruction by rearing pseudogynes. 
The latter modification of the nursing instinct, which 
leads to the rearing of cripples, can only be a 
pathological symptom, pointing to a morbid dis- 
turbance of the normal, organic condition of that 


1) “Die psychischen Faehigkeiten der Ameisen,” p, 111; ‘“In- 
stinct and Intelligence in the Animal Kingdom,” Chap. 8. 








Fig. 1. 


Form. san- 
guinea Ltr., 
subsp. 7udz- 
cunda Em. 
Normal dea- 
lated queen 
from Prairie 
du Chien, 
Wisconsin, 


U.S.A. 


























Fig. 2. 


Form, san- 
guinea Lir., 
subsp. ~udz- 
cunda Em. 
Normal wor- 
ker, from the 


same __ nest. 




















Fig. 3. 
Formica san- 
guinea Lirs 


subsp. rudbicun- 
da Em, Pseudo- 


gyne from the 


same nest. 


Le) 

















4 


(All figures magnified to seven times their natural size.) 


Fig. 4. 


Xenodusa cava 
Lec., whose lar- 
val 
caused the rea- 
ring of Pseudo- 
gynes in 
same colony of 


education 


the 


ants. 


Care of the Young in the Animal Kingdom. 181 


instinct. Hence, the improved education of the 
Lomechusa-larvae which is invariably attended by an 
impaired education of their own larvae, is not owing 
to intelligence on the part of Formica sanguinea, 
but to a disturbance of their normal, instinctive dispo- 
sition, occasioned by nursing the strangers. 

What we have said here of the rearing of our 
European Lomechusa strumosa by the sanguinea, has 
its exact counterpart in North America, where the 
larvae of Xenodusa cava, a species nearly allied to 
Lomechusa, are educated by the North American race 
of F. sanguinea, which Emery has named F. rubi- 
cunda. Rev. H. Muckermann, S. J., of Prairie du 
Chien (Wisconsin) has succeeded in finding also the 
pseudogynous ant-form in an infested nest of F. rubi- 
cunda. In the Verhandlungen der Deutschen Zoo- 
logischen Gesellschaft(1902 p. 98-108) I have given 
an account of these observations, which are illustrated 
on Plate II of the “Verhandlungen.” We give here 
a copy of this plate, to show more clearly what pseu- 
dogynes (fig. 3) are, and how they differ from the 
normal queen (fig. 2) and the normal worker (fig. 1) 
of the same ant-species. The malefactor, Xenodusa 
cava, is photographed in fig. 4. 

To be sure, at the first glance the care bestowed 
by ants on other animal species, their guests or nest- 
mates, often looks like intelligence. This explains to 
a certain extent, why modern animal psychologists 
attempted to utilize these occurrences as arguments 
for the great intelligence of ants. This attempt was 

1) Of late Father Muckermann himself has published an illustrated 


account of his discovery in the ‘Entomological News,” (Philadelphia), 
December, 1904. 


182 Chapter IV. 


made, e. g., by William Marshall in his “Leben und 
Treiben der Ameisen” (p. 102), where he speaks of 
the care given by ants to the eggs of plantlice. Sev- 
eral ant-species of the genus Lasius collect the eggs 
of certain Aphides in their nests. Being carefully 
protected during winter, the young aphides in spring 
are carried to the plants on which they find their food. 
Thence Marshall infers, that the ants tend the eggs 
with the intelligent purpose of enjoying later on the 
sweet secretions of the aphides. “This is surely a 
very strange phenomenon,” he says, “which proves 
perhaps better than anything else the high degree of 
intelligence attained by ants. We must credit them 
with a considerable power of observation, and we must 
own, that they have studied, to a certain degree, the 
habits of their domesticated animals,” etc. Yet this 
bold conclusion is entirely unfounded. How does 
Marshall know, that the ants gather the eggs of the 
aphides with the intelligent purpose of rearing 
aphides? That.there is some connection between the 
eggs of the aphides and the aphides themselves is, 
indeed, for many ants a subject of sensitive knowl- 
edge and experience; but it is unwarrantable to mis- 
take this process of instinctive association for intelli- 
gence proper. Even if ants in reality tended the eggs 
of aphides only on account of a combination of their 
sensitive experiences, this would be as yet no proof 
of their intelligence, but merely of their memory. 
In reality, however, the case is different. Take a few 
newly developed workers of a Lasius nest and unite 
them to form an autodidactic colony, restricted to its 
innate instincts without a shadow of experimental 


Care of the Young in the Animal Kingdom. 183 


knowledge as to the development of aphides. Entrust 
them with eggs of those aphis-species, which their 
congeners are wont to rear and to nurse. They will 
treat them as though they had previously “studied” 
the habits of these aphides! Hence the fondness of 
certain ant-species for the eggs of aphides is a merely 
instinctive impulse, which, of course, can be strength- 
ened by sensitive experience. It was rather rash for 
Mr. Marshall to proclaim it boldly as “a faculty of 
taking the future into account.’ Alfred Espinas was 
far more correct in calling the aphis-nursing of ants 
an “intelligence non réfléchie,” i. e., merely analogous 
to human reason, having but a faint similarity to 
intelligence proper, the difference being not merely of _ 
degree but of kind.t| This analogum rationis is simply 
an instinctive association of representations, assisted 
by-sensitive experience. 

In spite of the perfection attained in their nursing 
of plantlice, the Lasius species are far inferior to the 
Formica species in what modern animal psychology 
erroneously styles intelligence, viz: in the ability to 
profit for the future by past experiences. It will be 
interesting, therefore, to examine, in how far the latter 
ant-species, in taking care of their offspring, “con- 
sciously foresee the future.” 

Whenever care is taken of the young, then also the 
future is instinctively taken into account, above all 
in the rearing of the female ant-larvae; for it 
depends entirely on modifications in the nursing, 
whether the fertilized egg will produce a female 
proper or a worker. But only uncritical popular 


1) “Sociétés animales” (2d ed.), pp. 157, 188, etc, 


184 Chapter IV. 


psychology is able to confuse instinctive intention and 
instinctive foresight with intelligent intention and 
intelligent foresight. This is made evident by the 
following facts. The beetles of the genus Atemeles 
have their larvae reared by certain Formica-species, 
Atemeles emarginatus by F. fusca, Atemeles paradoxus 
by F. rufibarbis, Atemeles pubicollis by F. rufa, Ate- 
meles pratensoides by F. pratensis. The young Ate- 
meles having successfully reached their full develop- 
ment, either quit the Formica nests or are driven out of 
them. They then move over to Myrmica rubra’ and 
spend the greater part of their lives in the nests of 
these ants, by whom they are licked and fed. Only in 
spring, in the mating season, they return to their 
respective Formica species, where they allow their 
offspring to be reared at the expense of the ant-larvae. 
For whom, then, are these Formica species nursing 
the young Atemeles? Not for themselves, but for 
the Myrmica species. The only consequence of their 
adopting the Atemeles-larvae is the immense damage 
inflicted on their own eggs and larvae by these vora- 
cious changelings. Where now is “the faculty of 
intelligently taking the future into account,’ with 
which Marshall credits his ants? For thousands of 
years the Formica again and again have had the sad 
experience, that the pains bestowed on these beetle- 
larvae are but “love’s labor lost.” I believe that if 


1) This older collective name comprises Myrmica scarbrinodis, 
laevinodis, ruginodis, sulcinodis and rugulosa. 

2) The same applies to the education of the larvae of the North 
American Xenodusa in the nests of Formica species; for, the Xenodusa 
are found as fullgrown beetles with other ants, especially of the genus 
Camponotus, 





Care of the Young in the Animal Kingdom. 185 


Mr. Marshall had not been ignorant of the develop- 
ment of these myrmecophilous beetles, he would 
scarcely have extolled the great intelligence of ants. 

Lomechusa strumosa spends her whole life with F. 
sanguinea which, no doubt, is the “most intelligent” 
of European ants. She remains with her hosts after 
having attained the state of imago, and even, as a 
general rule, abides in the same nest, in which she 
was reared. In this case, therefore, the ants really 
derive some profit from rearing the Lomechusa-larvae. 
They not only have the pleasure of gratifying their 
nursing instincts by the rapid growth of these adopted 
children, but also later on they enjoy an agreeable, 
narcotic stimulant obtained by licking the yellow 
hair-tufts of the beetle. But if the ants had the 
faintest trace of intelligence, would they really be 
foolish enough to rear the Lomechusa-larvae merely 
for the sake of this sensual gratification? They 
experience again and again, that these changelings 
are their worst enemies, that they destroy their brood 
and moreover cause the birth of the merest cripples, 
namely the pseudogyne workers. Hence ants ought 
to have perceived long ago, that by rearing the Lome- 
chusa-larvae they are guilty of a folly little short of 
suicide. But alas, the very contrary is the case. The 
longer Lomechusa-rearing has been going on in a 
given sanguinea-colony—which time can be ascertained 
by the increasing number of pseudogynes,—the more 
care is bestowed on these beetles; and the percentage 
of beetle-larvae, which, after being imbedded in the 
earth, are left undisturbed, is continually increasing. 
The experiences made only serve to entangle the ants 


186 : Chapter IV.. 


more and more in the toils of their treacherous guests. 
In the face of such facts “animal intelligence” is alto- 
gether untenable. On the other hand, these facts 
furnish a new argument proving the correctness of 
our explanation of the psychic activities of animals. 

Birds which nurse the unfledged cuckoos, do not 
behave a whit more reasonably than the ants with 
regard to their Lomechusa-larvae. Because the young 
cuckoo opens its bill wider, makes more noise and 
wiggles its stumpy wings more energetically, its 
“foster-parents” feed it with special devotedness, and 
rather suffer their own young to starve. Moreover 
they calmly look on, whilst the young cuckoo pushes 
their own offspring over the edge of the nest to make 
them fall to the ground; indeed it has been observed, 
that the foster-parents assist in this work.1 Among 
birds, too, the nursing and adopting instincts are due 
to the very same laws of sensitive life as in ants. 
There is no discrimination between their own off- 
spring and that of others, no idea of “consanguinity,” 
of “parents” or “children,” but everywhere we wit- 
ness the same unreasoning dependence on instinctive 
sense-impressions, the appropriateness of which for 
the welfare of their own or of strange species escapes 
the sensitive knowledge of the animal. 

This is manifest also in the care bestowed on their 
young by the highest mammals, the apes. Just as 
within the same species of ants eggs, larvae, and 
pupae are a kind of international property, and are 
therefore received and nursed also by other colonies; 
as the eggs of eider ducks, of hens and other birds 


2) “Westfalens Thierleben,” II, 22, 


Care of the Young in the Animal Kingdom. 187 


have the same international character, extending even 
to the rearing of the young developed therefrom; as 
in many ants and birds the instinct of adoption, which 
is founded on the external resemblance between the 
nurslings of strangers and their own, is now and then 
extended to entirely different species (Lomechusa, 
cuckoo): so there is in apes a similar instinct owing 
to the same psychological causes, which proves to 
evidence the lack of intelligence in animals. “It is a 
well-known fact,” as the third ed. of Brehm’s Tierleben 
(p. 52) has it, “that apes, without much ado, adopt 
the children of any other species, protect them with 
the utmost tenderness, and can scarcely be separated 
from their dead bodies. When our shepherd-dog 
Trina would present us again with young puppies 
swarming with fleas, we used to put them into a cage 
of marmosets. There they were heartily welcomed, 
cleaned and fondled with care and tenderness, whilst 
from without the old dog was watching with a know- 
ing look (sic). But as soon as we deprived them 
of their nurslings, the monkeys would set up a pitiable 
screaming: they had distributed the pups among 
their number and evidently intended (sic) to keep 
them.” The anthropomorphism, with which modern 
fanatics in the matter of animal intelligence try to 
varnish over the true character of these adoption phe- 
nomena, must be mercilessly exposed by genuine, crit- 
ical psychology. We wish to picture the psychic life 
of the animals such as it is in itself, and not as it 
exists in the imagination of would-be psychologists. 

That the inclination of apes to adopt the offspring 
of other apes, of dogs, cats, rabbits, Guinea-pigs, and 


188 C. hapter IV. 


even of man is an entirely instinctive impulse devoid 
of intelligent reflection, is so evident to logical minds 
and so plainly expressed in the facts, that further 
proof seems superfluous. Since with apes the two 
sexes differ far less in psychic endowments than with 
ants, it can hardly be surprising, that not only the 
females but also the males have an instinctive nursing 
inclination, and try to gratify it by nursing any young 
animal. But how do they do it, especially if the young 
ones belong to another species? Alfred Brehm says,* 
“Here the ape often appears to be an inexplicable 
puzzle. He nurses his adopted favorite to the full 
extent of his power, hugs him, cleans him, continually 
keeps an eye on him, but generally does not supply 
him with any food. Without pangs of conscience 
(sic), he keeps for himself the food destined for his 
nursling, and even carefully keeps him away from the 
pot, whilst he himself is eating. This I have observed 
with baboons, who had picked up young dogs or cats 
as their foster-children.” 

Is this really an “inexplicable puzzle’? Only for 
those who are unwilling to understand the correct 
solution, because they are blinded by their monomania 
on animal intelligence. The solution of the puzzle is 
as clear as day-light. The instincts both of nursing 
and of eating are purely sensitive inclinations, unat- 
tended by reason and reflection. The faculties of 
sensitive cognition and appetite are so appropriately 
disposed in animals, that with regard to their own 
offspring the nursing instinct is stronger than hunger, 
but only so long as the young of that species, under 


1) Ibidem, p. 51. 


Care of the Young in the Animal Kingdom. 189 


normal circumstances, require nursing. Encaged she- 
apes often dispute every bite with their offspring, 
although they themselves have enough to eat; yea, 
they would even allow their “darling children” to 
starve, unless they were prevented by force, or unless 
the young possessed the strength and agility to get 
at the food in spite of the envy and greediness of their 
“dear mamma.” 

Critical psychology cannot but regard these phe- 
nomena as evident proofs, that even the highest 
animals are unreasonable beings, whose actions are 
guided only by instinctive impulses. Under normal 
conditions, the nursing instinct which serves to pre- 
serve the species, is, in animals, with regard to their 
own young, stronger than hunger, which provides for 
the preservation of the individual; the higher law of 
preserving the species demands so. Hence, in the 
beginning the mother-ape suckles and feeds her young 
with unrivalled “unselfishness,” whereas later on she 
grudges it every bite; hence apes hug and nurse the 
young of strangers with every sign of affection, 
whilst at the same time they deny them food and 
cruelly allow them to starve; hence the worker-ants 
nurse with motherly tenderness the eggs not laid by 
themselves, whereas for the most part they devour 
the eggs, which they have laid by way of partheno- 
genesis. Their natural duties being those of nurses 
and not of mothers, these loving aunts then become 
cannibals and monsters of. cruelty, because their 
instinct of eating is subordinated to that of nursing 
not by intelligence or consciousness of duty, but by 
the appropriate disposition of their sensitive appetite. 


190 Chapter IV. 


Let us briefly sum up the results of our discus- 
sion on the nursing instinct of animals. In this respect 
all animals obey the same psychological laws. Every- 
where the inclination of nursing and rearing the young 
proves to be a sensitive instinct, entirely different from, 
and even excluding, individual reflection and conscious- 
ness of duty. This is the case both in the highest 
mammals and in ants; for the latter even far surpass 
the highest mammals by their quasi-intelligent freedom 
of choice in rearing the different castes, and by an 
attachment to their charges verging on heroic unsel- 
fishness. With all animals the care of the young is 
directed exclusively by sensitive impulses and percep- 
tions, which, under normal circumstances, are suitably 
regulated both for preserving their own species and 
for maintaining the equilibrium between different 
species. Yet this appropriate correlation is far beyond 
the ken of the animal; hence, in the nursing of ani- 
mals there is no question of any “consciousness of 
duty.” Man alone by virtue of his intellect perceives 
the relations of consanguinity and the connections 
resulting therefrom; he alone has an intellectual 
notion of “parents” and “children”; only with him 
can there be question of the moral duties of parents 
toward their children. True, also in man motherly 
love is founded on a sensitive instinct; but, at the 
same time, it is spiritual, because the mother knows 
that she is the mother of this child, and because this 
knowledge with the resultant consciousness of the 
duty of attending to the welfare of the child, lasts for 
life. In man the love of parents toward their chil- 
dren and the care they bestow on them rises far above 


‘Care of the Young in the Animal Kingdom. 191 


the sphere of sensitive instinct into the province of 
spirituality and morality: and because the love of a 
mother is a rational love, conscious of duty, there- 
fore it is the highest and noblest love existing in 
nature. To ascribe such motherly love to animals, 
as do modern psychologists, is nonsense, scientifically 
speaking, and morally speaking it is a degradation of 
human dignity. 


CONCLUSION. 


OT to exceed the limits of this publication, we 

must refrain from pointing out other parallels 
existing between the psychic life of ants and that of 
the other animals. In particular, the extremely vari- 
ous ways of gaining a living, prevalent in ant-com- 
munities, would furnish plenty of material. But in 
this essay we had to confine ourselves to a few stray 
remarks on that subject (p. 38). What we have 
dilated upon may suffice, however, to furnish a posi- 
tive and reliable answer to the question, with which 
we introduced our essay, namely: Are animals en- 
dowed with instinct only, or also with intelligence? 

We have already proved in a former publication 
(Instinct and Intelligence in the Animal Kingdom), 
_of which the present study is a confirmation, that 
modern animal psychology influenced by so-called 
popular psychology, has inverted and confused the 
notions of sensitive cognition and of intelligence. 
That which. is popularly styled animal intelligence, 
in as far as it is based on real facts and not on fables 
and anecdotes, is nothing but the faculty of the ani- 
mals of forming complex representations of their 
Sensitive experiences and of acting appropriately in 
accordance with them. But this power as well as the 
immediate instinctive cognition is due to the innate 
laws of associations of sensitive representations and 
affections; hence it belongs to the sphere of sensitive 

192 


Conclusion. 198 


instinct, not to spiritual intelligence. By a critical, 
psychological analysis we were led to define instinct 
as the appropriate disposition of the sensitive powers 
of cognition and appetite. Hence, any action result- 
ing therefrom must be called instinctive, whether 
experience is concerned in it or not. But only those 
actions can be called intelligent, which presuppose the 
understanding of the relations existing between the 
sensitive representations, and which cannot be ex- 
plained by any other supposition. Intelligence, there- 
fore, exclusively signifies the power to act with deliber- 
ation and self-consciousness. Only this power can be 
called a spiritual faculty; the sensitive power of repre- 
sentation and memory cannot be so called, notwith- 
standing the efforts of modern animal psychologists. 
The pretended “spiritual life’ of animals, about which 
popular psychology continues to make such ado, is 
based on the confusion between sensitive and spiritual 
faculties. 

Modern animal psychology splits up the psychic 
life of animals into two different factors, styled 
instinct and intelligence, between which an artificial 
contrast is established. Our explanation of the psychic 
life of animals is more consistent and more natural. 
What is erroneously termed animal intelligence, we 
have traced to the same source as_ the instinctive 
actions strictly so called, namely, to the suitable 
hereditary disposition of the sensitive cognition and 
‘appetite, which we call “instinct;’ for, this dispo- 
sition has a twofold aspect, one automatic, the other 
plastic. It is automatic, inasmuch as it is determined 
by ata and therefore induces the animal to per- 


\ 


194 Conclusion. 


form certain actions, which are independent of indi- 
vidual experience and are more or less the same in 
all individuals of a given species. It is plastic, inas- 
much as within this limited sphere, the powers of 
cognition and appetite in the animal are given more 
or less play for variously modifying their activities. 
The narrower the limits within which they are con- 
fined, the more automatically their instinct will cause 
them to act; the wider those limits, the more plastic 
their instinct. Both elements, automatism and plas- 
ticity, are found in different proportions -with all 
animals from the highest to the lowest. In the lower 
orders automatism, as a general rule, largely prevails, 
whereas in the higher vertebrates plasticity is, on the 
average, more predominant. Ants, too, more than 
dogs and apes, are bound by hereditary laws to the 
performance of certain activities. The varying 
influence, which individual sensation brings to bear 
upon the performance of hereditary instincts, is greater 
and more variable in the latter than in the former, 
and in this respect the psychic life of ants is more like 
“automatism” than that of mammals. But, on the 
other hand, the plasticity of the instinct is, also in 
ants, often highly developed, and not rarely it is 
manifested in a more quasi-intelligent form, than even 
in the highest vertebrates. 

In the present essay we have reviewed a number 
of the most prominent phenomena of the psychic life 
of animals, and everywhere we found, that, what 
modern animal psychology styles animal intelligence, 
is met with also in ants and in many cases, in fact, in 
a higher degree than with the highest mammals. In 


Conclusion. 195 


the community-life of ants, which with suitable 
co-operation for the welfare of the colony combines 
a manifold independence of action on the part of the 
single workers, in their mutual communications and 
mutual services, in their wars, in their slave-making 
expeditions and their confederations, in their nest 
construction and in the manifold application of their 
building skill to various changes of circumstances, 
finally, in their breeding and nursing, embracing 
various methods of education left to the choice of the 
workers and manifesting, at the same time, the highest 
degree of “self-sacrificing attachment” to their help- 
less young ones: in all these points combined we must 
rightly consider the life of ants as the climax of 
development in imstinctive life throughout the animal 
kingdom. As regards the perfection of the nervous 
system and of the sense-organs, the higher mammals 
are indeed far closer to man, than the ants; but as 
regards the quasi-intelligent actuation of animal 
instinct under the influence of sense-perceptions and 
experiences for the various purposes of community- 
life, ants no doubt approach nearer to man than even 
the anthropoid apes. Indeed, neither of them pos- 
sesses intelligence proper, that is to say, the power 
of acting with deliberation and self-consciousness, of 
inventing new means for attaining various purposes 
and thus making progress in civilization. Still, the 
chasm between the psychic life of animals and that 
of man, is, in many respects, wider between ape and 
man, than between ant and man. 

Of course, the results of our study are very dif- 
ferent from, and indeed altogether contrary to, the 


196 Conclusion, 


aprioristic postulates of modern evolutionism,? accord- 
ing to which man is nothing but the highest brute, 


1) We cannot enter here on the general question of the develop- 
ment of instincts. Cf. for this purpose my former publications: “Die 
Entstehung der Instincte nach Darwin” (“‘Stimmen aus Maria-Laach,” 
XXVIII, 333), “Die Entwicklung der Instincte in der Urwelt” (ibid. 
papi 481; XXIX, 248, 383); “Zur Entwickl hichte der 

lischaften” (‘‘Die zusammengesetzten Nester und gemischten 
Kolosien der Ameisen,” III section, 2 Chap.). 

As to the explanation of the genuine guest-relationship (symphily) 
by the Darwinian theory of evolution cf. “Zur Entwicklung der In- 
stincte” (“Verhandlgn. der Zoolog. Botan. Gesellsch.,” Wien, 1897, 3d 
issue, pp. 168-183). Of late Dr. K. Escherich has tried to solve the 
contradiction, which we proved to exist between the facts of 
symphily and the principles of natural selection (“Zur Anatomie und 
Biologie von Paussus turcicus, Zugleich ein Beitrag zur Kenntnis der 
Myrmekophilie,” in ‘‘Zoolog. Jahrbuecher” Abth. fuer Systematik, XII, 
1898, 27-70). He insists that symphily is not a separate instinct 
totally different from the breeding instinct of ants, but that the two are 
in causal relation to each other. We formerly (see the first German ed. 
of present essay, p. 107 below, and p. 108 above) pointed out the same. 
Yet Escherich is wrong in believing that natural selection has been 
unable to prevent the development of symphily in spite of the damage 
done by it to the ants, because symphily is so closely connected with 
the breeding instinct. Natural selection must counteract not only the 
development of an entirely new instinct which proves injurious to the 
possessor, but also the extension to injurious objects of an already 
existing useful instinct; hence selection was just as little allowed to let 
the breeding instinct of ants extend jits activity to Lomechusa, Atemeles, 
Paussus and other noxious objects, as it was allowed to let the feeding 
instinct of animals extend its activity to palatable but poisonous herbs 
or to nutritious plants covered with parasites. (Cf. “‘Die psychischen 
Fachigkeiten der Ameisen,” 1899, p. 124.) To this Escherich again 
objected (in “‘Zool. Centralbl.,” 1899, No. 1, p. 17), that many sheep 
are killed by feeding on plants covered by “cercaries” (i. ¢., the 
capsulate form of undeveloped trematodes). But what would Mr. 
Escherich say to the following, if within the whole species of sheep, 
or within a certain race of them, there should develop a special liking 
‘for feeding on plants covered by those parasites? Would not such a 
phenomenon evidently contradict the theory of natural selection? But 
this is exactly the case with the rearing of Lomechusas by the 
sanguine slaveé-makers. Therefore Escherich’s objections but confirm 
the truth of our assertion: The fact that ants by nursing their guests 
rear their greatest enemies, ts equally incompatible with the principles 
of natural selection and with the principles of modern animal psychology. 








Conclusion, 197 


and human society but a gradual evolution from that 
of the higher mammals. But scientific research can- 
not be hampered by such aprioristic theories; if they. 
are incompatible with facts, they are to be abandoned. 
It is an undeniable fact, that between the soul of man 
and that of the brute there yawns a chasm, which 
cannot be bridged over by any evolutionistic specula- 
tion.1 Man is, as a matter of fact, the only being in 
the visible universe, who is gifted with reason, with 
a spiritual soul, and with morality. On account of 
the essential difference between sensitive and spiritual 
life, it is simply impossible, that in the course of 
nature an animal should ever develop into man. True, 
we can daily witness, how from instinctive sensations 
children gradually arrive at spiritual reasoning; but 
this development is possible only because from the 
outset the soul of the child is a sensitivo-spiritual soul. 
The development of its spiritual faculties must be 
preceded by sensitive instincts, because these furnish 
the foundation and the materials for the spiritual 
faculties. The animal, however, which never mani- 
fests spiritual faculties, cannot be credited with any- 
thing beyond a sensitive soul, which is essentially dif- 
ferent from the sensitivo-spiritual soul of man, and 
which makes the animal, be it ape or ant, a being 
devoid of reason, and belonging to a lower order! 
Hence, so-called popular animal psychology, which 
denies the essential difference between the human 
spirit and the animal soul, and which appeals in favor 


1) Even evolutionists like Wallace have well understood this, and 
therefore they protest against applying Darwinism to the psychic part 
of man. Cf. Wallace, “Darwinism.” 


198 Conclusion, 


of this theory to the results of biological research, 
must im the first place, be branded as unscientific ;+ 
for it mistakes sensation for spiritual life, and 
instinct for intelligence, thus being diametrically at 
variance with the principles of critical psychology. 
Secondly, its assertion, that the brute is gifted with 
reason and consciousness of duty as well as man, 
although in a different degree, is an evident falsehood, 
which is given the lie by the actual biological facts. 
But this popular psychology is not only unscientific 
and untruthful; it is far worse. To be candid, it is 
demoralizing and fraught with moral danger to the 
human social order. Hence we must do more than 
merely shrug our shoulders in contemptuous pity, we 
must take a decided stand against it and combat it 
with all our might. 

By denying the existence of the essential differ- 
ence between animal and human psychic faculties, this 
psychology not only raises the brute to the level of 
man, but degrades man to the level of the brute. 
Would to God that this were done in theory only; 
but, alas, the practical consequence of this false theory 
is the demoralization and brutalization of man. This 
is the goal aimed at by those books and pamphlets, 


1) Let me once more protest, as I have already done in Chap. I of 
“Instinct and Intelligence in the Animal Kingdom,” that there is no 
wish on my part to identify the scientific representatives of our modern 
zoological psychology with the champions of animal intelligence like 
Brehm, Buechner, etc. This would be an injustice to very many 
sober-minded naturalists, who condemn just as we do the humanization 
of the animal. Nor do we in any manner intend to pass judgment on 
the personal motives of Brehm, Buechner and other defenders of 
' animal intelligence and animal morality, but we only judge of their 
writings. This remark is added here expressly to avoid misunder- 
standings. 


Conclusion, 199 


which describe the sexual impulse of the brute as 
essentially the same as human conjugal love, and the 
care of the young among animals as essentially iden- 
tical with parental love in man. Such men as Alf. 
Brehm and L. Buechner were not ashamed to come 
forward as “apostles of free love” and to decry as 
antiquated and ridiculous the moral bounds estab- 
lished for man by reason and divine law. With them 
the humanization of the brute, consciously or uncdn- 
sciously, aims at degrading man so far as to make 
him cast off his reasonable nature and to follow with- 
out reserve the sensual inclinations, which he has in 
common with the unreasonable brute. On this account 
they deny the difference between sensitive and spirit- 
ual faculties, between the animal soul and the human 
spirit. Hence we do not consider it too harsh a judg- 
ment to say: Those, who humanize the animal, not 
only trifle with scientific psychology, but they also 
drag into the mire the dignity of man. Every well- 
meaning naturalist, therefore, ought resolutely to 
oppose these unprincipled doirgs of so-called popular 
psychology. 

Now-a-days, there is, and rightly so, a widespread 
agitation against the use of alcohol and other drugs 
injurious to the nervous system, because the bodily 
and spiritual welfare of humanity is endangered. But 
to counteract the ravages of spiritual venoms, which 
under the glittering name of modern science are spread 
through all classes of society, little or nothing is done. 
If the moral principles of Brehm and Buechner should 
later on become the common property of humanity, 
then the society of the future from the highest to the 


200 Conclusion, 


lowest, would resemble a herd of unreasonable ani- 
mals, whose “spiritual life’ would consist in the 
unbridled gratification of the meanest lusts and pas- 
sions. Hence our concluding appeal: Do away with 
all books, pamphlets and periodicals, whose only pur- 
pose is to raise the brute to the level of man! 








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